Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela

Conde, J. E.; Tognella, M. M. P.; Paes, E. T.; Soares, M. L. G.; Louro, I. A.; Schaeffer Novelli, Y. Population and life history features of the aratus pisonii (: grapsidae) in a subtropical estuary Interciencia, vol. 25, núm. 3, mayo-junio, 2000, pp. 151-158 Asociación Interciencia Caracas, Venezuela

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POPULATION AND LIFE HISTORY FEATURES OF THE CRAB ARATUS PISONII (DECAPODA: GRAPSIDAE) IN A SUBTROPICAL ESTUARY

J. E. Conde1, M. M. P. Tognella2, E. T. Paes2, M. L. G. Soares2, I. A. Louro2 and Y. Schaeffer-Novelli2 SUMMARY

Population features of the grapsid crab Aratus pisonii, a similar in the two study sites: 1.63 individuals/m2 in Bertioga-D common inhabitant of the supralittoral zone of roots, branches and 1.41 in Bertioga-Cidade. Females were significantly more and canopy of several of , were studied in abundant than males in a ratio of 1.168:1.00 in Bertioga-D, and two estuarine forests near Bertioga, São Paulo, Bra- 1.153:1.00 in Bertioga-Cidade. J-shaped sex ratio curves were zil. were larger in Bertioga-D (range: 8.85 - 26.20 mm), observed for both populations. Juvenile crabs were present at where a well-developed mangrove forest is present, than in both locations, but they were much more abundant in Bertioga- nearby Bertioga-Cidade (range: 3.80 - 20.85 mm), where Cidade (34.8%) than in Bertioga-D (3.9%), possibly as a conse- arbustive and stunted mangrove trees prevailed. Ovigerous fe- quence of the topography of each site. This study in a subtropi- males were present at both sites; the percentages in Bertioga-D cal estuary suggests that a conceptual model presented previ- and in Bertioga-Cidade were similar, 31.0% and 32.1%, respec- ously, relating population features of A. pisonii and structural tively; however, the mean number of eggs was larger in Bertio- characteristics of mangroves forests for various localities on the ga-D than in Cidade, 15,367 (maximum: 33,533) and 10,433, Venezuelan coast, could be extended. (maximum: 19,994) respectively. Abundances of A. pisonii were

Introduction tropical estuarine areas mangals Crabs are one of the most strate topography and man- are remarkable for sheltering a important groups, so grove community structure, Mangrove woodlands once protracted catalogue of inverte- much for their high diversity in population dynamics, and life occupied some 70% of tropical brate species, thus being the some regions, as for their im- histories (Malley, 1978; Beever coastlines. Mangroves play a main vegetation in supporting portance in the flows of energy et al., 1979; Lacerda, 1981; key role in maintaining coastal biodiversity (Twilley et al., in these forests and bushes. As Giddins et al., 1986; Robert- stability and water quality, as 1996). Although mangroves a consequence of their diversity son, 1986; Warren and Under- well as in promoting soil devel- have received extensive atten- and roles, a growing interest in wood, 1986; Smith, 1987a, opment (Twilley et al., 1995). tion during centuries, most of the ecology of mangrove crabs 1987b; Díaz and Conde, 1988, In some arid regions, man- the research efforts have been has been arising in the last 1989; Camilleri, 1989; Conde groves provide true coastal oasis aimed at vegetational aspects, years (Lee, 1998). Progress has and Díaz, 1989a, 1989b; Con- for non-mangrove species and while much remains to be been made in knowledge on de et al., 1989; Neilson and stepping stones in otherwise learned about most of the asso- their feeding behavior, nutrient Richards, 1989; Robertson and barren expanses. Furthermore, in ciated fauna. cycling role, impact on sub- Daniel, 1989; Smith et al.,

KEY WORDS / Mangrove productivity / Life history / Stunting / Reproductive Effort / Brazil /

1 Centro de Ecología, Instituto Postal 21827, Caracas 1020-A, 2 Instituto Oceanográfico, Uni- Venezolano de Investigaciones Venezuela. versidade de São Paulo, CEP Científicas (IVIC), Apartado E-mail: [email protected] 05508-900, São Paulo, Brazil.

MAY - JUN 2000, VOL. 25 Nº 3 0378-1844/00/03/151-08 $ 3.00/0 151 RESUMEN

En dos manglares estuarinos próximos a Bertioga (São cias de A. pisonii fueron similares en ambos sitios; 1,63 indivi- Paulo, Brasil) se estudiaron las características poblacionales del duos/m2 en Bertioga-D y 1,41 en Bertioga-Cidade. La proporción cangrejo Aratus pisonii, un grápsido muy común en la zona de sexos se inclinó significativamente hacia las hembras, tanto en supralitoral de las raíces, ramas y dosel de varias especies de Bertioga-D (1,168:1,00), como en Bertioga-Cidade (1,153:1,00). mangles. El tamaño de los cangrejos fue significativamente mayor Las curvas de proporción de sexos asumieron forma de J en en Bertioga-D (ámbito: 8,85 - 26,20 mm), en un bosque de man- ambas poblaciones. Probablemente debido a la topografía de los glar bien desarrollado, que en Bertioga-Cidade (ámbito: 3,80 - sitios de estudio, los juveniles de esta especie representaron 20,85 mm), en el que dominaban mangles arbustiformes y 34,8% en Bertioga-Cidade; mucho más que en Bertioga-D, donde achaparrados. En los dos sitios se encontraron hembras ovígeras apenas alcanzaron 3,9% del total de individuos recolectados. Este en proporciones similares: 31,0% en Bertioga-D y 32,1% en estudio, llevado a cabo en un estuario subtropical, sugiere que Bertioga-Cidade. Sin embargo, el número promedio de huevos puede extenderse un modelo conceptual que relaciona las varia- por hembra fue mayor en Bertioga-D (15.367; maximo: 33.533) ciones en poblaciones de A. pisonii con las características estruc- que en Bertioga-Cidade (10.433; máximo: 19.994). Las abundan- turales de los manglares de la costa venezolana.

RESUMO

Aspectos populacionais do caranguejo Grapsidae Aratus 1,63 individuos/m2 em Bertioga-D e 1,41 em Bertioga-Cidade. pisonii, habitante comum da zona supralitoral de raízes, troncos Fêmeas foram significativamente mais abundantes que os ma- e galhos de diversas espécies de mangues, foram estudados em chos com razões fêmeas/machos de 1,168:1,00 em Bertioga-D, e dois manguezais na região estuarina de Bertioga, São Paulo, 1,153:1,00 em Bertioga-Cidade. Curvas de taxas entre sexos do Brasil. Os indivíduos foram maiores em Bertioga-D (8,85 - tipo forma de J foram observadas nas duas populações. Juvenis 26,20 mm), caracterizada por um manguezal bem desenvolvido, estavam presentes nas duas áreas, mas muito mais abundantes que na área próxima à cidade Bertioga-Cidade (3,80 - 20,85 em Bertioga-Cidade (34,8%) que em Bertioga-D (3,9%), possi- mm), dominada por arbustos de mangue pouco desenvolvidos. velmente como consequência da topografia de cada área. Os re- Fêmeas ovígeras ocorreram nas duas áreas com percentuais si- sultados deste estudo em um estuario subtropical sugere que o milares (31,0% e 32,1% respectivamente). Entretanto, o número modelo conceitual previamente apresentado, relacionando carac- medio de ovos foi maior em Bertioga-D que em Bertioga-Cidade terísticas populacionais de A. pisonii e características estrutu- (15.367; máximo: 33.533 e 10.433; máximo: 19.994, respectiva- rais das florestas de manguezais de várias localidades da Vene- mente). A abundância de A. pisonii foi similar nas duas áreas: zuela, pode ser extendido para outras regiões.

1989; Wilson, 1989; Warren, coast (Garth, 1960; Chace and 1990; Lacerda et al., 1991; Mi- Hobbs, 1969; Beever et al., cheli et al., 1991; Emmerson 1979; Branco, 1991; Conde, and McGwynne, 1992; Miche- personal observations). A. li, 1993a, 1993b; Davie, 1994; pisonii has been found in a Frusher et al., 1994; McIvor wide spectrum of habitats: and Smith, 1995; McGuinness, river mouths, estuarine and 1997). But in spite of this en- marine habitats, and hypersa- larging body of information, it line lagoons, where it is asso- is considered that there is little ciated with several species of quantitative information on mangroves, including Rhizo- community structure and popu- phora mangle, Avicennia ger- lation dynamics (Twilley et al., minans, A. schaueriana, La- 1996). Particularly, the factors guncularia racemosa, and the that regulate populations of tea mangrove, Pelliceria rhizo- mangrove are still phorae. In some of these habi- poorly known. tats, Aratus has shown consid- Most of the information erable plasticity in body size available on population dy- and life history traits (Conde namics and life histories has and Díaz, 1989b, 1992a; Con- been provided by several en- de et al., 1989; Díaz and Con- tries dealing with the Neotro- de, 1989). In low productivity pical mangrove crab Aratus mangroves that grow in hyper- Figure 1. Map of the Brazilian coast showing the study sites in the pisonii (H. Milne Edwards saline lagoons in the western State of São Paulo. A: Bertioga-Cidade. B: Bertioga-D. 1837) (Brachyura: Grapsidae). coast of Venezuela, popula- This crab is one of the few tions of undersized individuals brachyurans with an amphi- of A. pisonii, including unusu- mangroves growing in river Aratus, such as maximum American distribution, ranging ally small ovigerous females, mouths (Conde, 1989; Conde body size, reproductive effort, from eastern and most were observed (Conde et al., and Díaz, 1992b). Based on and sex ratios, to mangrove of the Antilles to Santa 1989; Conde and Díaz, these relations, Conde and productivity, as inferred from Catarina (Brazil) on the Atlan- 1992b); while the largest indi- Díaz (1989b) sketched a con- forests structural characteristics tic coast; and from northern viduals and highest reproduc- ceptual model that relates a (Conde and Díaz, 1989b). Mexico to on the Pacific tive effort were recorded in series of population features of However, most of the knowl-

152 MAY - JUN 2000, VOL. 25 Nº 3 edge about the biology, life ties at the Canal da Bertioga, dominant species (94.7% base females distribution in that site. history and population features a channel located east of area and 77.5% of total indi- Recruitment patterns, growth of A. pisonii springs from Santos, State of São Paulo, viduals), followed by Rhizo- rate and longevity were esti- tropical populations located in Brazil (23° 50´ S, 46° 05´ W) phora mangle and Avicennia mated by means of the the Caribbean (Hartnoll, 1965; (Figure 1). The channel is schaueriana, which have a ELEFAN (Electronic Length Hagen, 1977; Warner, 1967; connected to the open sea by basal area of, respectively, 1.8 Frequency Analysis) computer Díaz and Conde, 1988, 1989; means of two wide inlets, and 3.1; and 9.3 and 13.1% of package (Brey and Pauly, Conde, 1989; Conde and Díaz, which allow expeditious water total individuals (Peria et al., 1986). This method can be 1989a, 1989b, 1992a, 1992b; exchange. Mean annual tem- 1990). used with a single sample Conde et al., 1989). The ex- perature in the area is 21.3°C (Pauly and David, 1981) pro- ceptions in this corpus are the (range: 6.4 - 39.0°C) and the Materials and Methods vided that the sample is large works of Lacerda (1981) and average annual precipitation is enough and representative of a Lacerda et al. (1991), who 2,080.8mm (Santos, 1965; Crabs were captured accord- major sector of the population have examined the feeding Ministério da Agricultura e ing to the catch-effort design during a stable phase. It does habits of Aratus in Brazil; Reforma Agrária, 1992). Since described by Conde and Díaz require the simplifying assump- Branco (1991), who reported there is a continuous input of (1989a). At each site, in an tion, as with the determination abundances of this crab in fresh water from several rivers, area of 50m2, bounded by of population parameters from southern Brazil; and Arruda the waters of the channel are means of cords (10 x 5m), any single age frequency Leme and Negreiros-Fransozo estuarine, ranging from 21 to three persons caught all man- sample, that these are more or (1998), who studied reproduc- 34%. The surface salinity of grove crabs in sight on the less similar each year; thus, to tive patterns. the mangrove swamps water branches, aerial roots and in this extent, ELEFAN provides Being a highly plastic spe- ranges from 8 to 35, whereas the underwater portion of sub- a first, although useful, ap- cies that has shown substantial the interstitial water fluctuates merged roots of mangrove proximation. Data were com- ecological differences on local between 14 and 28 (Schaeffer- trees during half an hour. The pared by means of standard and regional scales, and even Novelli, unpublished data). cephalothorax width (CW) of statistical tests (t-test, G-test, in populations separated by a Tides follow a semidiurnal each crab (i.e. size) was meas- etc.; Sokal and Rohlf, 1995). few hundred meters, it is inter- pattern with a maximum of ured by means of Vernier cali- Except in those cases where a esting to determine if similar 1.6 m over the mean level, pers with a precision of 0.05 confidence interval has been differences hold for warm which is 0.7m (Brasil, 1988). mm. Sex and presence of eggs specifically indicated, the vari- temperate populations that are The site designated as Ber- were also assessed. After- ability is expressed in terms of very close to its Atlantic tioga-D is located approxi- wards, most of the crabs were ± the standard error. southern distributional limit mately 17.5 km northeast of released alive at sites of col- and thus are exposed to a dif- Santos, between the Caiubura lection. Each 50m2 plot was Results ferent set of environmental and Itapanhaú rivers (Figure considered a replicate. Four conditions. In this investiga- 1). This mature mangrove for- replicates were obtained in Population Structure and tion we seeked to widen the est, some 200m from the Bertioga-D, while in Bertioga- Abundance available knowledge on the channel, has a mean density of Cidade two replicates were population features of Aratus 256 individuals/0.1 ha and a carried out, due to time con- A total of 107 males and pisonii, and therefore of man- basal area of 1.13m2/0.1 ha. straints. Samplings were un- 125 females of Aratus pisonii grove crabs, by addresing sev- The mean height and diameter dertaken on April 13, 15 and were collected and measured eral topics regarding the popu- at breast height (dbh) of man- 17, 1994. in Bertioga-D, while in Bertio- lation characteristics of this groves in this location are, re- Ovigerous females were col- ga-Cidade a total of 70 males crab in an estuary. Our objec- spectively, 7.7m (± 0.7) and lected in areas close to the and 83 females were caught tives were: 1. To determine if 7.5cm (Peria et al., 1990). The sampling sites, and preserved and measured. In Bertioga-D, the body dimensions of A. dominant species is Rhizo- individually in 10% formalin in the carapace width (CW) fre- pisonii in a subtropical estuary phora mangle with 36.6% of seawater. In the laboratory, the quency distributions of both show the same patterns ob- basal area; Avicennia schaueri- number of eggs in the berry males and females were bell- served in populations that in- ana and Laguncularia race- was estimated by means of the shaped (Figure 2), whereas in habit arbustive and arboreal mosa follow with 28.6 and volumetric method described Bertioga-Cidade the CW fre- mangroves in hypersaline la- 16.7%, respectively, while by Díaz et al. (1983). Males quency distributions of males goons in the Caribbean; 2. To dead trees represent 18% of and non-ovigerous females and females were polymodal gather information about popu- cover (Peria et al., 1990). The were put in an icebox and (Figure 3) and at least three lation features of A. pisonii second location, herein desig- taken to the laboratory, where peaks can be distinguished for close to its distributional limit nated as Bertioga-Cidade, is their carapace width and both sexes. In both sites, the in the South Atlantic; and 3. located next to the town of length, and length and width of shapes of males and females To determine if body sizes of Bertioga, approximately 22.0 the left chelae were recorded. frequency distributions differed this brachyuran are larger in km northeast of Santos, near Size at the onset of maturity significantly (Bertioga- D: higher latitudes, a common re- the eastern inlet of the chan- (SOM) was determined follow- G = 13,932, P < 0.01, d.f. = 4; lationship in crustaceans. nel. This forest, contiguous to ing the criterion in Conde and Bertioga-Cidade: G = 42,608, the channel, has a mean den- Díaz (1992a); that is, SOM P < 0.01, d.f. = 4). Study Area sity of 963 individuals/0.1 ha, was considered as the smallest Crabs were larger at Bertio- a basal area of 0.48 m2/0.1 ha, of two numbers: the smallest ga-D. The percentage of crabs Sampling was taken in two mean height of mangroves is ovigerous female collected in a with CW larger than 21.00mm mangrove forests, selected be- 3.1 m (± 0.3) and their dbh is given site, or the inferior limit was 15.9% at this site, where- cause of their clear-cut struc- 2.5cm (Peria et al., 1990). La- of the 99% confidence interval as in Bertioga-Cidade none of tural differences, at two locali- guncularia racemosa is the for the mean of the ovigerous the crabs surpassed this cutoff

MAY - JUN 2000, VOL. 25 Nº 3 153 recruitment in Cidade was spread evenly through the year, with 69.5% of the re- cruits arriving from June to October, peaking in Septem- ber. In Bertioga-D, however, estimated recruitment was more confined: 67.2% oc- curred in just three months, from April to June (maximum in May, 28%).

Reproductive Effort

As indicated by the pres- ence of ovigerous females, re- productive activity was ob- served in both localities. Dur- ing the half-hour sampling pe- riods, 36 ovigerous females were randomly collected in Bertioga-D; while 17 were caught in Bertioga-Cidade. These numbers were supple- mented with 36 and 42 ovig- Figure 2. Carapace width frequency distributions Figure 3. Carapace width frequency distributions erous females randomly caught of males and females of Aratus pisonii from Ber- of males and females of Aratus pisonii from Ber- in the surrounding areas of tioga-D. tioga-Cidade. Bertioga-D and Bertioga-Ci- dade, respectively. The per- value. In Bertioga-D males males CW ranged from 3.80 from the normal Mendelian ra- centage of ovigerous females CW ranged from 9.45 to to 19.70 mm. tio (1.00:1.00). In Bertioga-D in the two locations, 31.0% in 26.20mm, while females CW Crabs were equally abun- there were 1.168 females for Bertioga-D and 32.1% in Ber- ranged from 8.85 to 24.35 dant at both sites (t = 1.065; every male in the population tioga-Cidade, did not differ mm. The average CW size of 0.4 > P > 0.2; d.f. = 4). In Ber- (G = 1.398; 0.5 > P > 0.1; d.f. significantly (G = 0.024, 0.9 > males (18.54 ± 0.325mm) was tioga-D the mean abundance = 1; G test), while this quo- P > 0.5, d.f. = 1). Carapace significantly larger (t = 2.580, was 1.63 individuals/m2, while tient was 1.153 in Bertioga- width of ovigerous females 0.01 > P > 0.001) than that of in Bertioga-Cidade the density Cidade (G = 0.781; 0.5 > P ranged from 12.45 to 23.50 − females (17.43 ± 0.286mm). was slightly, but nonsignifi- > 0.1; d.f. = 1). Sex ratios did mm in Bertioga-D (χ=17.79 ± However, in Bertioga-Cidade cantly lower (1.41). not differ significantly between 0.268); while in Bertioga-Ci- the mean CW of males (13.48 both locations (G = 0.0041; dade ovigerous females were ± 0.520mm) did not deviate Sex Ratio and 0.975 > P > 0.9; d.f. = 1). significantly smaller (t = significantly (t = 1.272, 0.4 Sex Ratio Curve Sex ratio curves were J- 3.991, P < 0.001; d.f. = 129), >P> 0.2) from that of females shaped (Figure 4), resembling ranging from 11.95 to 21.90 − (12.61 ± 0.450mm). At this lo- At both sites the sex ratios the “standard pattern” of mm (χ =16.54 ± 0.271). The cation, males CW ranged from of A. pisonii were female-bi- Wenner (1972). These curves CW distributions differed sig- 5.90 to 20.85 mm, while fe- ased and deviated significantly show that males predominated nificantly (G = 8.216; 0.01 in the largest size classes, while < P < 0.05; d.f. = 3). The dif- females accumulated in the ferences in reproductive effort middle and lower size classes. can also be assessed by exam- In several instances the 95% ining the proportions of oviger- confidence intervals are large ous females that were larger or due to low sample size in equal to 19.00 mm; in Bertio- those particular segments. ga-D it was almost one-fourth (23.6%), while in Bertioga- Recruitment Cidade this proportion was substantially smaller (10.2%). Juvenile crabs (CW < 10 In both sites the number of mm; Conde and Díaz 1989a) eggs per female increased as a were present in both popula- linear function of CW (Figure tions, but their percentage var- 5). Both relationships were sta- ied significantly (G = 67,328; tistically significant (Bertioga- P < 0.001; d.f. = 1). The high- Cidade: r = 0.651, P < 0.01; est proportion of recruits was Bertioga-D: r = 0.801, P < 0.01). observed in Bertioga-Cidade For the 42 values regressed in Figure 4. Sex ratio curves of Aratus pisonii from Bertioga-Cidade (34.8%), while in Bertioga-D Bertioga-Cidade, fecundity and Bertioga-D. it was only 3.9%. Estimated ranged from 4,816 to 19,994

154 MAY - JUN 2000, VOL. 25 Nº 3 eggs, with an average of 10,433 eggs/female. In Bertio- ga-D, for a given size, females carried more eggs, with a mean number of 15,367, and a range of 5,634-33,533.

Maturity, Longevity and Relative Growth

In Bertioga-Cidade crabs were estimated to attain matu- rity within 2.5 years, while the maturational process was faster in Bertioga-D, where maturity should be reached in 1 year. Size at the onset of maturity (SOM) confirms these trends; SOM was higher (10.97mm) for females from Bertioga-D than for females from Bertio- ga-Cidade (10.29). Longevity of crabs, as estimated through ELEFAN, was highest in Ber- tioga-D than in Bertioga- Cidade; 4.5 and 6 years, re- spectively. For each population the Figure 5. Scattergram of the relationship between number of eggs and carapace width in equations that describe the females of Aratus pisonii from Bertioga-Cidade and Bertioga-D. growth of chelae relative to the carapace width are pre- suggests that salinity, although served by Micheli (1993a) The ultimate factors respon- sented in Table I. According probably important, is not the whose data, based on labora- sible for these relationships, to the form of equations, rela- only factor modulating life tory experiments with the however have not been identi- tive growth for females in histories in this crab. Thus, it mangrove crab Sesarma fied, although several possi- Bertioga-D and Bertioga-Ci- was interesting to determine if messa, hint that there is a di- bilities have been scrutinized dade were very similar. How- the predictions of the concep- rect link between mangrove (Conde et al., 1989; Conde ever, the chelae of males in tual model would hold in an forest primary production and and Díaz, 1992b). Growth re- Bertioga-Cidade were rela- estuary with forests showing secondary production of crabs. tardation in aquatic inverte- tively larger than those of different degrees of structural In other marine environments, brates, resulting in individuals males from Bertioga-D. development. Based on the body dimensions of organisms smaller than those of normal present study, conducted in a have also been related to pro- size, has been attributed to a Discussion subtropical estuary in which ductivity. For instance, Ansell number of factors including body dimensions and fecun- and Lagardere (1980) ascribed salinity, food depletion, para- Conde and Díaz (1989b) re- dity differences were found, diverging maximum sizes sites, dissication, and other ported correlations between the model can be extended. In among populations of bivalves stressful physical conditions several population features of this estuary, specimens of A. to differences in productivity (Newman and Pollock, 1974; Aratus pisonii, including body pisonii were larger on arboreal among habitats. Thus, the va- Chittleborough, 1976; Brown size and reproductive effort, mangroves and their fecundity, lidity of this conceptual rela- and Caputi, 1985; Hawkes et and structural characteristics of as compared with crabs of tionship seems well-founded al., 1986). Not enough is mangrove forests for various similar sizes from arbustive and merits further reckoning in known about Aratus pisonii localities on the coast of mangroves, was also higher. other mangrove crabs and also to distinguish between those Venezuela. The largest Some of these relationships in other wide-ranging man- mechanisms; however, since and the highest reproductive are similar to the ones ob- grove invertebrates. mangrove productivity and effort were observed in a population of A. pisonii living in riverine mangroves, while TABLE I the smallest crabs and the RELATIVE GROWTH EQUATIONS OF MALES AND FEMALES OF ARATUS PISONII lowest reproductive effort were FROM BERTIOGA-CIDADE AND BERTIOGA-D. CHL: observed in hypersaline la- CHELA LENGTH. CW: CARAPACE WIDTH. goons. However, even in these lagoons large crabs were observed in arboreal Site Males Females mangroves, as compared to crabs inhabiting arbustive Bertioga-Cidade ChL = -2.9428 + 0.7307 CW ChL = -0.9558 + 0.4899 CW mangroves in the same la- goon; a result that strongly Bertioga-D ChL = 0.1388 CW 1.485 ChL = -1.5277 + 0.5316 CW

MAY - JUN 2000, VOL. 25 Nº 3 155 structural development can be 1989a). In Bertioga-D, 15.9% were similar, clutch size was this population trait is most determined by several ele- of the specimens of Aratus larger in Bertioga-D, where fe- likely a result of the topo- ments or conditions (Lugo pisonii exceeded 20.95 mm, males carried up to 33,533 graphic profile of the man- and Snedaker, 1974; Rodrí- while in Laguna de Tacarigua eggs, while in Bertioga-Cidade grove forest rather than the guez, 1987; Saenger and this quotient was much higher the maximum number of eggs productivity of the environ- Snedaker, 1993), these should (42.4%). Additionally, the in a berry was 19,994. These ment. be examined to shed some maximum size of Aratus in differences arise from the larg- The results of this study light on growth retardation that lagoon was 29.45 mm, est size of ovigerous females provide insights into several and life history variations in compared to 26.20 mm in in Bertioga-D, but there is also population characteristics of mangrove crabs. Also, various Bertioga-D, so there appears an effect that goes beyond the mangrove crab Aratus structural characteristics of to be a reduction of crab sizes morphological constraints and pisonii. However, we are mangrove forests are related that could be attributed to lati- could be due to environmental aware of the obvious limita- to their productivity (Scha- tude. In like manner, varia- restrictions, as can be noted tions that stem from the re- effer-Novelli and Cintrón, tions in size at the onset of when the overlapping sections strictions in the number of 1986) and consequently to the maturity (SOM) supports a of ovigerous females sizes are sampling dates. Even though stressors to which these plants latitudinal effect. Size at the compared. The regression recruitment and longevity can are exposed. Likely, these onset of maturity (SOM) for equation that describes the re- be estimated by means of the characteristics also influence A. pisonii in Venezuelan estu- lation between female carapace ELEFAN package when only life history characteristics of aries and river mouths ranged width and number of eggs in one or two sampling dates are the Neotropical grapsid A. from 13.35 to 15.68 mm Aratus pisonii collected in available, aliasing could arise pisonii. Salinity has been (Conde and Díaz, 1992a), Bertioga-D is similar to that in these cases, leading in turn commonly regarded as the while at both sites of Bertioga found in mangrove trees in an to the concealment of the real key element in growth retar- this parameter was below estuary on the Venezuelan magnitude of fluctuations. dation in crustaceans; how- 11.00 mm. However, further coast, and so are the mini- Thus, conclusions regarding ever, it does not seem to play data collection is advisable be- mum, maximum and average these particular parameters are a definitive role in this spe- fore jumping to conclusive number of eggs (Conde and to be qualified and deemed cies, since salinities are simi- generalizations. Díaz, 1989a); however, the re- cautiously. Likewise, this lar in both study sites. Thus, Besides cephalothorax width, gression parameters of females study does not pretend to pre- other agents should be in- other morphometric differences from Bertioga-Cidade, an es- empt much needed long-term voked to explain growth re- were recorded. The chelae of tuarine population, are closer population dynamics studies duction and life history varia- males caught at Bertioga- to the scope of a hypersaline comprising monthly sam- tions in Aratus. Furthermore, Cidade were relatively longer population in the Venezuelan plings throughout several Frusher et al. (1994) found than those of males from Ber- coast. These facts support the years. Still, the values ob- that four species of mangrove tioga-D. Relative growth for perception that the effect of tained for most of the param- crabs belonging to the females from both sites were the salinity on the physiology eters furnish information that Sesarma have a wide toler- similar. The significance of of Aratus is not the crucial el- widens the understanding of ance to salinity, ranging from these differences is not fully ement in the population fea- the relations between man- completely fresh to hypersa- understood, but could be re- tures shown by this crab and grove crabs and their habitats. line waters, and that salinity lated to the biomechanical that variation should also be tolerance did not explain their properties of mangrove leaves. ascribed to other variables. ACKNOWLEDGEMENTS distributions. The findings of In two Brazilian mangrove Overall, the estimated re- this study and the fact that forests, where A. pisonii was cruitment intensity and its pat- This work was possible marine-sized ovigerous fe- present, Lacerda et al. (1986) terns through time in both lo- thanks to the generous finan- males were found on arboreal found that crude fiber and ash cations were unlike. Few re- cial support of the U.S. Fish mangroves in a hypersaline contents in mangrove leaves cruits were present in Bertio- and Wildlife Service and the lagoon in Venezuela, while were negatively correlated to ga-D and most of the esti- Academia de Ciencias do smaller crabs were present on herbivory. On the Venezuelan mated recruitment should be Estado do São Paulo (Brazil). arbustive mangroves in the coast, arboreal mangroves concentrated in a few months, Laboratory facilities and same lagoon (Conde et al., growing in estuaries and river as estimated through ELE- transportation were provided 1989; Conde, 1989), suggest mouths are less coriaceous and FAN; whereas recruitment in by the Instituto Oceanográfico the same conclusions. not so heavily grazed as the Bertioga-Cidade was higher de la Universidade de São A latitudinal explanation to leaves of arbustive and stunted and appears to be spread Paulo. Special thanks are the differences observed in mangroves (JE Conde and C through the year. The physiog- given to Bibiana Molina and Bertioga obviously is not vi- Alarcón, unpublished data). raphy of both sites might ex- Henry Ramos (IVIC) for their able since the populations Crabs inhabiting these man- plain such differences: the assistance in fecundity estima- sampled are only a few km groves could be selected for mangrove forest of Bertioga- tions. Our gratitude is ex- apart. However, there seems to more rugged chelae to respond Cidade borders the channel, so tended to Arno Blankensteyn, be a latitudinal effect over to the increased difficulties of it can receive larvae of Aratus Paulo Cesar de Paiva, María body dimensions, when the scrapping more coriaceous continuously, while the man- Helena de Arruda Leme, crabs collected in the arboreal leaves. grove forest in Bertioga-D is Valter J. Cobo, Ricardo San- mangroves of Bertioga-D are In addition to the somatic approximately 200 m inland tos da Silva and Andrea S. compared to those caught in dissimilarities, there were also and it is connected to the Freire for valuable help dur- the arboreal mangroves of La- differences in the reproductive channel by a shallow water- ing data gathering. Luiz Dru- guna de Tacarigua, a Carib- effort at both sites. Although way that would allow the en- de de Lacerda kindly wrote bean estuary on the Venezu- the proportions of ovigerous trance of recruits only during the Portuguese version of the elan coast (Conde and Díaz, females at the two localities exceptionally high tides. Thus, abstract.

156 MAY - JUN 2000, VOL. 25 Nº 3 REFERENCES Conde JE, Díaz H (1989b) Giddins RL, Lucas JS, Neilson rine Biology and Ecology Productividad del habitat e MJ, Richards GN (1986) 171: 149-163. Ansell AD, Lagardere F (1980) historias de vida del cangrejo Feeding ecology of the man- Micheli F (1993b) Feeding ecology of Observation on the biology of de mangle Aratus pisonii (H grove crab Neosarmatium mangrove crabs in North Eastern Donax trunculus and D. Milne-Edwards) (Brachyura: smithi (Crustacea: Decapoda: Australia: mangrove litter con- vittatus at Ile D’Oleron Grapsidae). Boletín del Ins- ). Marine Ecology sumption by Sesarma messa. and (French Atlantic Coast). Ma- tituto Oceanográfico de Vene- Progress Series 33: 147-155. Sesarma smithii. Journal of Ex- zuela (Universidad de Orien- rine Biology 57: 287-300. Hagen HO von (1977) The tree- perimental Marine Biology and te) 28: 113-120. Arruda Leme MH, Negreiros- climbing crabs of Trinidad. Ecology 171: 165-186. Fransozo ML (1998) Repro- Conde JE, Díaz H (1992a) Varia- Studies on the Fauna of Micheli F, Gherardi F, Vannini M ductive patterns of Aratus tions in intraspecific relative Curaçao, and Other Carib- (1991) Feeding and burrowing pisonii (Decapoda: Grapsidae) size at onset of maturity bean Islands 54: 25-59. ecology of two east African (RSOM) in Aratus pisonii (H from an estuarine area of São Hartnoll RG (1965) Notes on the mangrove crabs. Marine Biol- Milne Edwards, 1837) (Deca- Paulo Northern Coast, Brazil. marine grapsid crabs of Ja- ogy 111: 247-254. poda, Brachyura, Grapsidae). Revista de Biología Tropical maica. Proceedings of the Crustaceana 62: 214-216. Ministério da Agricultura e Reforma 46: 673 - 678. Linnean Society of London Agrária (1992) Normais clima- Beever III JW, Simberloff D, King Conde JE, Díaz H (1992b) Exten- 176: 113-417. tológicas (1961-1990). Minis- LL (1979) Herbivory and pre- sion of the stunting range in Hawkes CR, Meyers TR, Shirley tério da Agricultura e Reforma dation by the mangrove tree ovigerous females of the man- TC (1986) Length-Weight re- Agrária, Brasília, Brasil. crab Aratus pisonii. Oecologia grove crab Aratus pisonii (H. lationships of blue and golden Neilson MJ, Richards GN (1989) 43: 317-328. Milne Edwards, 1837) (De- king crabs parasitized by the Chemical composition of de- capoda, Brachyura, Grap- Branco JO (1991) Aspectos eco- rhizocephalan, Briarosaccus grading mangrove leaf litter sidae). Crustaceana 62: 319- lógicos dos Brachyura (Crus- callosus Boschma. Fishery and changes produced after 322. tacea: Decapoda) no mangue- Bulletin 84: 327-332. consumption by the mangrove zal do Itacorubi, SC—Brasil. Conde JE, Díaz H, Rodríguez G crab Neosarmatium smithi Lacerda LD (1981) Mangrove Revista Brasileira de Zoología (1989) Crecimiento reducido (Crustacea: Decapoda: Sesar- wood pulp, an alternative food 7: 165-179. en el cangrejo de mangle Ara- midae). Journal of Chemical source for the tree-crab Aratus tus pisonii (H Milne-Edwards) Ecology 15: 1267-1283. Brasil (1988) Tábua das marés pisonii. Biotropica 13: 317. para o ano de 1988, costa do (Brachyura: Grapsidae). Acta Newman GG, Pollock DE (1974) Brasil e portos estrangeiros. Científica Venezolana 40: 159- Lacerda LD, Jose DV, Rezende Growth of the rock lobster DHN. Ministério da Marinha, 160. CE, Francisco MCF, Wasser- Jasus lalandii and its relation- man JC, Martins JC (1986) Brasil. Davie PJF (1994) Variations in di- ship to benthos. Marine Biol- Leaf chemical characteristics Brey T, Pauly D (1986) Electronic versity of mangrove crabs in ogy 24: 339-346. affecting herbivory in a New Length Frequency Analysis. A tropical Australia. Memoirs of World mangrove forest. Bio- Pauly D, David N (1981) ELEFAN revised and expanded user’s the Queensland Museum 36: tropica 18: 350-355. 1, a BASIC program for objec- guide to Elefan 0, I and II. 55-58. tive extraction of growth pa- Institute fur Meereskunde, Lacerda LD, Silva CAR, Rezende Díaz H, Conde JE (1988) On the rameters from length-frequency Kiel. 76 pp. CE, Martinelli LA (1991) food sources for the mangrove data. Meerforschung/Reports Food sources for the man- Brown RS, Caputi N (1985) Fac- tree crab Aratus pisonii on Marine Research 28: 205- grove tree crab Aratus pisonii: tors affecting the growth of (Brachyura: Grapsidae). Bio- 211. a carbon isotopic study. undersize western rock lobster, tropica 20: 348-350. Peria LCS, Fernandes PPCP, Revista Brasileira de Biología Panulirus cygnus George, re- Menezes GV, Grasso M, Tog- turned by fishermen to the sea. Díaz H, Conde JE (1989) Popula- 51: 685-687. tion dynamics and life history nella MMP (1990) Estudos es- Fishery Bulletin 83: 567-574. Lee SY (1998) Ecological role of of the mangrove crab Aratus truturais comparativos entre grapsid crabs in mangrove eco- Camilleri J (1989) Leaf choice by pisonii (Brachyura: Grapsidae) bosques de mangue impactados systems: a review. Marine and crustaceans in a mangrove in a marine environment. Bul- (Canal da Bertioga) e não Freshwater Research 49: 335- forest in Queensland. Marine letin of Marine Science 45: impactados (Ilha do Cardoso), 343. Biology 102: 453-459. 148-163. Estado de São Paulo. In: II Chace Jr FA, Hobbs HH (1969) Lugo AE, Snedaker SC (1974) Simpósio de Ecossistemas da Díaz H, JE Conde, Bevilacqua M The freshwater and terrestrial The ecology of mangroves. Costa Sul e Sudeste Brasileira: (1983) A volumetric method decapod crustaceans of the Annual Review of Ecology Estrutura, Função e Manejo. for estimating fecundity in West Indies with special refer- and Systematics 5: 39-64. Aguas de Lindóia, SP. 6 a 11 ence to Dominica. United Decapoda. Marine Ecology de abril de 1990. Publicação Progress Series 10: 203-206. Malley DF (1978) Degradation of States National Museum Bul- ACIESP Academia de Ciências mangrove leaf litter by the letin 292: 1-258. Emmerson WD, McGwynne LE do Estado de São Paulo Nº 71- tropical sesarmid crab Chiro- (1992) Feeding and assimila- 2 Vol. 2: 183-193. Chittleborough RG (1976) Growth manthes onychophorum. Ma- tion of mangrove leaves by the Robertson AI (1986) Leaf-burying of juvenile Panulirus longipes rine Biology 49: 377-386. cygnus George on coastal crab Sesarma meinerti de Man crabs: their influence on en- reefs compared with those in relation to leaf-litter produc- McGuinness KA (1997) Seed preda- ergy flow and export from reared under optimal environ- tion in Mgazana, a warm-tem- tion in a tropical mangrove for- mixed mangrove forests (Rhi- mental conditions. Australian perature south African man- est: a test of the dominance- zophora spp.) in northeastern Journal of Marine and Fresh- grove swamp. Journal of Ex- model in northern Australia. Journal of Experi- water Research 27: 279-295. perimental Marine Biology and Australia. Journal of Tropical mental Marine Biology and Ecology 157: 41-53. Ecology 13: 293-302. Ecology 102: 237-248. Conde JE (1989) Ecología pobla- cional del cangrejo de mangle Frusher SD, Giddins RL, Smith McIvor CC, Smith III TJ (1995) Robertson AI, Daniel PA (1989) Aratus pisonii (H Milne-Edw- III TJ (1994) Distribution and Differences in the crab fauna of The influence of crabs on lit- ards) (Brachyura: Grapsidae) abundance of grapsid crabs mangrove areas at a Southwest ter processing in high inter- en hábitats extremos. Ph. Sc. (Grapsidae) in a mangrove Florida and a Northeast Austra- tidal mangrove in tropical Dissertation, Instituto estuary: effects of sediment lia location: implications for Australia. Oecologia 78: 191- Venezolano de Investigaciones characteristics, salinity toler- leaf litter processing. Estuaries 198. Científicas, Caracas. ances, and osmoregulatory 18: 591-597. Rodriguez G (1987) Structure and Conde JE, Díaz H (1989a) The ability. Estuaries 17: 647- Micheli F (1993a) Effect of man- production in neotropical man- 654. mangrove tree crab Aratus grove litter species and avail- groves. Trends in Ecology and pisonii in a tropical estuarine Garth J (1960) Distribution and ability on survival, moulting, Evolution 2: 264-267. coastal lagoon. Estuarine, affinities of the brachyuran and reproduction of the man- Saenger P, Snedaker SC (1993) Coastal and Shelf Science 28: Crustacea. Systematic Zoology grove crab Sesarma messa. Pantropical trends in mangrove 639-650. 9: 105-123. Journal of Experimental Ma- above-ground biomass and an-

MAY - JUN 2000, VOL. 25 Nº 3 157 nual litterfall. Oecologia 96: Smith III TJ (1987b) Seed preda- system function: ecosystem tidal predators of temperate 293-299. tion in relation to tree domi- analysys. Cambridge Univer- mangrove crabs. Marine Santos E de O (1965) Características nance and distribution in man- sity Press, UK. 387 - 393. Ecology Progress Series 67: climáticas. In: A De Azevedo grove forests. Ecology 68: 266- Twilley RR, Snedaker SC, Yáñez- 295-299. (Ed.). Baixada santista: aspectos 273. Arancibia A, Medina E (1996) Warren JH, Underwood AJ (1986) geograficos. EDUSP, São Paulo, Smith III TJ, Chan HT, McIvor Biodiversity and ecosystem pro- Effects of burrowing crabs on Brasil. Cap 4. pp. 95-150. CC, Robblee MB (1989) Com- cesses in tropical estuaries: Per- the topography of mangrove Schaeffer-Novelli Y, Cintrón G parison of seed predation in spectives of mangrove ecosys- swamps in New South Wales. (1986) Guia para estudo de tropical tidal forests from three tems. In: Mooney HA, Cush- Journal of Experimental Ma- áreas de manguezal; estrutura, continents. Ecology 70: 146- man JH, Medina E, Sala OE, rine Biology and Ecology função e flora. Caribbean Eco- 151. Schulze E-D (Eds.). Functional 102: 223-235. logical Research, São Paulo, Sokal RR, Rohlf FJ (1995) Biom- roles of biodiversity: a global Wenner AM (1972) Sex ratio as Brasil. 150 pp. + 3 appendixes. etry. 3d. ed. Freeman WH, San perspective. John Wiley & a function of size in marine Sons. 327 - 370. Smith III TJ (1987a) Effects of Francisco, USA. xix + 887 pp. Crustacea. The American seed predators and light level Twilley RR, Snedaker SC, Yáñez- Warner GF (1967) The life history Naturalist 106: 321-350. on the distribution of Avicen- Arancibia A, Medina E (1995) of the mangrove crab, Aratus Wilson KA (1989) Ecology of nia marina (Forsk.) Vierh. in Mangrove systems. In: Hey- pisoni. Journal of Zoology mangrove crabs: predation, tropical, tidal forests. Estua- wood VH (Ed.). Global biodi- (London) 153: 321-335. physical factors and refuges. rine, Coastal and Shelf Science versity assessment. Functional Warren JH (1990) Role of bur- Bulletin of Marine Science 44: 25: 43-51. roles of biodiversity and eco- rows as refuges from sub- 263-273.

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