Population structure and reproductive period of the tropical hermit crab Calcinus tibicen (Decapoda : Diogenidae) in the region of Ubatuba, São Paulo, Brazil

The population of the hermit crab Calcinus tibicen in the Ubatuba region was studied with emphasis on seasonal abundance, seasonal size-frequency distribution, sex-ratio, and reproductive period based on the percentage of ovigerous females. The animals were collected at 2-month intervals for 2 consecutive years (from January 1993 to November 1994). Total mean animal size in shield length was 5.14 +/- 1.23 mm for males, 4.23 +/- 0.79 mm for females, and 4.53 +/- 0.60 for ovigerous females. A discontinuity in reproduction was observed, with an absence of ovigerous females in July (winter) and a high incidence from September to May (spring to autumn). Calcinus tibicen is sexually dimorphic in relation to size, with a higher growth rare in males. The sex-ratio pattern changes with season, indicating that it is more a function of season or habitat than of size.

Ecological distribution and reproductive period of the hermit crab Loxopagurus loxochelis (Anomura, Diogenidae) on the northern coast of São Paulo State, Brazil

The present study aimed to determine the ecological distribution of Loxopagurus loxochelis as a function of selected environmental factors, as well as its reproductive period based on the combined analysis of the presence of ovigerous females and the development of the gonad. The collections were carried out monthly from January to December 2000 in the region of Ubatuba (SP), using a fishing boat equipped with two double rig nets, in isobaths of 5, 10, 15, 20, 25, 30, 35 and 40 m, at which depths samples of sediment and water were collected. The cephalothoracic shield length and sex of the animals were determined; the abdomen was dissected to verify gonad development stages. A total of 334 individuals was obtained ( 196 males, 48 ovigerous females and 90 non-ovigerous females) with sizes varying from 3.0 to 8.6 mm (5.7 +/- 1.0 mm). A greater abundance of L. loxochelis (95.2%) was observed at depths of 20 and 25 m. These sites mainly revealed a substratum stable with a very fine sand fraction, which facilitates the habit of embedding shown by this hermit. With regard to gonadal analysis, it was possible to classify four gonad development stages for each sex: immature, rudimentary, developing and developed. It was observed that in the winter months about 71% of the females had gonads either developing or developed and 67% were ovigerous. Therefore, the winter period can be considered the peak of reproduction for this species, characterizing a typical seasonal-continuous reproduction.

Juvenile development and growth patterns in the mud crab Eurytium limosum (Say, 1818) (Decapoda, Brachyura, Xanthidae) under laboratory conditions

The post-larval development of the mud crab Eurytium limosum was studied under laboratory conditions by using the offspring of ovigerous females collected at the Comprido River mangrove, SP, Brazil. The first crab stage is fully described and the juvenile development, until crab stage 10, is examined with emphasis on morphological change, sexual differentiation and growth patterns. The carapace of the first crab stage is nearly square as observed in other xanthids, becoming similar to adults only at stage 15. The sexes can be distinguished from stage four, based on the number of pleopods and their morphology. While the intermoult period increases, the moult percentage decreases at each stage. The abdominal allometric growth is sex-dependent, with males showing a negative (b=0.71) and females an isometric (b=0.95) relative growth pattern. Male gonopods undergo a positive allometric growth, and their shape changes remarkably until sexual maturity. The cheliped dentition can be observed after stage 4. Regardless of sex, most crabs have a molariform right cheliped, which is thought to aid the handling of asymmetric prey such as gastropods.

Growth rate of the crab Dissodactylus crinitichelis Moreira, 1901 (Crustacea : Decapoda : Pinnotheroidea) under laboratory conditions

The goal of the present study was to investigate the growth rate of the crab Dissodactylus crinitiehelis, its molt increments, and the duration of the intermolt intervals under laboratory conditions, focusing on differences between sexes and between juvenile and adult phases. Crabs were collected at Flamengo Beach, Ubatuba, São Paulo State, Brazil, by scuba divers. In the laboratory, individuals were maintained in isolation and fed nauplii of Artemia sp. daily. The sex and carapace width of exuviae and dead crabs were recorded. During the juvenile phase, mean growth was 11.8 +/- 2.7% in males and 11.2 +/- 3.6% in females, with their respective intermolt intervals 25.4 +/- 9.2 and 26.4 +/- 8.3 d (mean +/- SD). After reaching morphological sexual maturity, the intermolt intervals increased to 33.4 10.1 d in males and 32.7 +/- 10.4 d in females, and the growth rates of both sexes slowed to 6.4 +/- 1.9% in males and 5.7 +/- 1.6% in females. There was a significant decrease in molt increment and an increase in intermolt intervals associated with the maturation from juveniles to adults, evidencing differential energy allocation during the last phase of ontogeny.

Morphometric study of the ghost crab Ocypode quadrata (Fabricius, 1887) (brachyura, ocypodidae) from Ubatuba, São Paulo, Brazil

A morphometric study of the ghost crab Ocypode quadrata from Ubatuba, São Paulo, Brazil, focusing on the relationships between carapace width and carapace length, abdomen width, gonopod length and propodus length and height of the major cheliped is presented. An allometric positive growth was found for most of the analyzed relationships; however, important changes in the gonopod and abdomen were observed at the size at which sexual maturity is attained. Morphological (external) sexual maturity for both sexes is achieved at a smaller size than that at which gonad maturity is reached. Concerning heterochely, 53.3% of the analyzed specimens exhibited a longer propodus length in the right cheliped.

Size variation in the grapsid crab Aratus pisonii (H. Milne-Edwards, 1837) among populations of different subtropical mangroves

Size variation in the adult population of the grapsid crab Aratus pisonii (H. Milne-Edwards, 1837) was studied by comparing median values obtained from six different populations from nearby mangrove areas. The mangrove structure at those sites was also examined and related to size differences among populations. Disregarding sex, crab size is apparently related to the mangrove structure. Smallest and largest crab sizes were recorded at the sites where tree density and tree diameter were minimum and maximum, respectively. Estimates of size at the onset of maturity, considering the size of the smallest ovigerous female, followed the same trend. These results support the hypothesis that average crab size and size at the onset of maturity can be associated to mangrove productivity.

On the population biology of the mottled shore crab Pachygrapsus transversus (Gibbes, 1850) (Brachyura, Grapsidae) in a subtropical area

The mottled shore crab P. transversus is probably the most common brachyuran crab living in the marine rocky intertidal of southeastern Brazil. However, its biology is largely unknown. In this study, some aspects of the population biology of this species are investigated. Distribution of individuals in the intertidal range is heterogeneous. Sabellariid worm reefs and mytilid mussel beds can be considered as nursery grounds retaining most part of juveniles, while the rocky surface is mostly inhabited by adults, which find safety shelters in rock crevices. Annual size frequency distributions revealed clues of population stability and indicated that young individuals reach the adult habitat in a gradual process. Sex-ratio follows the 1:1 proportion in smaller size classes but is biased towards males in larger ones. Higher mortality in larger females is indicated as a possible cause of this pattern. Ovigerous ratio shows a remarkable seasonality in which highest percentages of ovigerous females occur during summer months, while recruitment of young is more intense from April to July, suggesting a very extensive larval development. Once established in the intertidal zone, young recruits will develop to mature individuals in late spring, when molting crabs become scarce and proportion of breeding females increases. Therefore, growth and reproduction are in a great part temporally separated, allowing an annual life cycle in which settlement, growth and breeding may take place within a I-yr period.

Allometry of the secondary sexual characters of the shore crab Pachygrapsus transversus (Gibbes, 1850) (Brachyura, Grapsidae)

The allometric growth of secondary sexual characters in Pachygrapsus transversus is investigated from the 2(nd) crab stage onward. Clear sexual dimorphism is restricted to abdominal morphology, but ANCOVA analyses showed that chelae become larger in males and the carapace becomes wider in females. Size at the puberty moult in both sexes was estimated using Somerton's computer techniques. Mature II analyses applied to bi-log gonopod length vs, carapace length relationships indicated a puberty moult at 5.0 mm in males.In females, Mature I analyses detected the overlapping growth phase lines in bi-log carapace length vs. abdomen width scatterplots. Fitting the logistic equation provided an estimate of 50% maturity at 5.5 mm. The regression lines separate young and resting individuals from the potentially reproductive females, but they do not separate young from adult crabs. Year-round monthly samples showed that the proportion of small adult-like females is higher during the breeding season. After breeding, females may moult to a young-like morphotype, as observed in controlled laboratory conditions. Moulting to a resting condition splits smaller mature females into different growth phase lines. Therefore, estimates of female size at sexual maturity by means of abdomen allometric growth analyses are inadequate in this species.

A morphometric study of the mud crab, Panopeus austrobesus Williams, 1983 (Decapoda, Brachyura) from a subtropical mangrove in South America

The relative growth of the xanthid crab, Panopeus austrobesus was investigated by means of the allometric method. Crabs were obtained in the mangrove formed by the estuary of the rivers Comprido and Escuro (23degrees29'24S 45degrees10'12W), Ubatuba, São Paulo State, Brazil. All crabs were measured to obtain their carapace width (CW) and length (CL), abdomen width (AW) at the basis of the 5(th) somite, and major cheliped propodus length (PL) and height (PH). Males were also measured for their gonopod length (GL). The size of crabs based on CW ranged from 4.0 to 44.8 mm for males and 3.1 to 34.5 mm for females. The relative growth equation (Y = aX(b)) based on the relationship between GL and CW suggested that males reach their sexual maturity near 14.6 mm CW. Such relationship shows a positive allometry during the juvenile phase and an isometric growth in adult life. In females, the estimated size at 50% maturity is 13.0 mm CW, based on the relationship AW vs. CW. Males reach larger sizes than females, which probably provides them better conditions to protect females during courtship. Concerning cheliped size, approximately 73% of the crabs analysed (N = 209), disregarding sex, have the right PL larger than the left. The PL growth shows that specimens with a left major cheliped (26%) have a higher allometric coefficient, despite being smaller considering their CW. Such a difference may compensate the smaller size of the crab during defense or prey capture.

Reproductive behavior of the swimming crab Arenaeus cribrarius (Lamarck, 1818) (Crustacea, Brachyura, Portunidae) in captivity

In this study, the reproductive behavior exhibited by Arenaeus cribrarius in captivity was described, and the duration of each behavioral stage was measured. Swimming crabs were trawled in Ubatuba, northern littoral of São Paulo State, Brazil, and maintained in aquaria. Water conditions and food items were provided according to this species' natural requirements in the wild. In the presence of premolt females, intermolt males exhibited a courtship display that became intensified when the potential mate was visually perceived. After mate selection, the male carried the female under itself (precopulatory position) for 29.8 +/- 5.1 d until the female molted. Afterwards, the male manipulated the recently molted female, and inverted her position under itself as to penetrate her with his first pair of pleopods (copulation), a process that took 17.1 +/- 4.6 h. After copulation the male continued to carry his soft-shelled mate for 29.7 +/- 5.8 d (postcopulatory position). The time elapsed between copulation and spawning was 57.8 +/- 3.8 d and the time interval between successive spawns 33.8 +/- 7.1 d. Total embryonic development took 13.5 +/- 2.1 d in temperature conditions of 25.0 +/- 2.0 degrees C. During the last 4.7 +/- 1.4 d embryos' eyes were already visible. The reproductive behavior pattern in A. cribrarius is very similar to those previously described in other portunids.

Reproduction of the speckled swimming crab Arenaeus cribrarius (Brachyura : portunidae) on the Brazilian coast near 23 degrees 30 ' S

The reproductive biology of Arenaeus cribrarius from Ubatuba, São Paulo State, Brazil, was studied. Swimming crabs were sampled monthly for two years with otter trawls in two bays. A total of 941 males and 1,012 females were examined. Mating took place mainly in autumn involving postmolt females and intermolt males. At that time, gonad regression was verified in adult males, due to spermatophore transfer, and the molting of adult females. Ovigerous females or females with mature gonads were present year-round but more frequently captured during spring and summer. We found that 19, of all adult females were premolt, which indicated the occurrence of another mature instar and thus the absence of a well-defined terminal molt after puberty. Intermolt males were captured throughout the whole study period.

Morphometry of the crab Hexapanopeus schmitti (Decapoda : Xanthoidea) on the northern coast of the state of São Paulo, Brazil

A morphometric study of the xanthoid crab Hexapanopeus schmitti was carried out, using the allometric method. Samples were taken monthly for two years (1998-1999) in the Ubatuba region, northern coast of São Paulo, Brazil. Sex and size were assessed for each specimen, and all crabs were measured to obtain their carapace width (CW) and length (CL), abdomen width (AW) of females, major cheliped propodus length and height (PL and PH), and gonopod length (GL) of males. A total of 301 crabs were analyzed, 209 males and 92 females. The CWs of the crabs ranged from 2.5 to 9.8 mm for males and from 2.8 to 9.4 mm for females. The relative growth equation (y=ax(b)) based on the relationship between GL and CW suggested that males reach their morphological sexual maturity near 6.1 mm CW. In females, the estimated size at 50 % maturity was 4.8 mm CW, based on the relationship of AW vs. CW. Males reach larger sizes than females, which probably favors their ability to guard the females during courtship. In approximately 83 % of the crabs (n= 371), disregarding sex, the right cheliped was larger.

Sexual maturity of the speckled swimming crab Arenaeus cribrarius (Lamarck, 1818) (Decapoda, Brachyura, Portunidae), in the Ubatuba Littoral, São Paulo state, Brazil

The sizes at morphological and physiological maturity of male and female Arenaeus cribrarius were estimated to determine if both events are synchronous. Animals were captured with otto-trawls at Ubatuba, Brazil. A total of 2356 specimens, 977 males and 1379 females, were obtained. The major carapace width without spines (CW), the propodus length of the major cheliped (PL) and the width of the 5th abdominal somite (AW) were measured with vernier calipers. Allometric relationships and gonadal development were analyzed to determine the maturity in both sexes. The size at the onset of male morphological maturity was estimated at CW 52 mm, smaller than the CW 63.4 mm physiological maturity size observed. For females, these events are synchronous since both estimates converged at CW 59.7 mm. The onset of functional sexual maturity in A. cribrarius at CW 63.4 and 59.7 mm in males and females, respectively, would indicate a minimum size of CW 64 mm for fishing purposes. Differences between allometric and gonadal estimates indicate the importance of considering both methods. A comparison of the present results with other available data in portunid crabs is provided.

Population structure and breeding season of the South Atlantic hermit crab, Loxopagurus loxochelis (Anomura, Diogenidae) from the Ubatuba region, Brazil

A population of Loxopagurus loxochelis was studied in terms of seasonal abundance, size frequency distribution, sex ratio, and reproductive period (percentage of ovigerous females). Specimens were collected monthly over a period of two years (from September 1995 to August 1997) in the non-consolidated areas of the Ubatuba, Mar Virado, and Ubatumirim bays (northern coast of São Paulo State, Brazil) with a double-rig trawl net. A total of 1,084 individuals were analysed. Animal size (minimum, maximum, and mean +/- SD shield length) was 2.8, 9.1, and 6.88 +/- 1.13 mm for 625 males; 2.8, 8.2, and 5.78 +/- 0.98 mm for 236 non-ovigerous females; and 4.6, 8.0, and 6.24 +/- 0.68 mm for 223 ovigerous females, respectively. Sexual dimorphism was recorded by the presence of males in the largest size classes. The sex ratio was 1.4 : 1 in favour of males. The highest incidence of ovigerous females occurred during winter and spring (June to October), with a low percentage in summer and fall (November to May), indicating continuity in the reproductive cycle. This strategy of reproduction is related to temperature. The occurrence of L. loxochelis in the Ubatuba region represents the final point of northern distribution of this species as a function of water mass influence, and could be its real limit on the Atlantic coast of South America.

Breeding season of the hermit crab petrochirus diogenes (anomura : diogenidae) in the north coast of São Paulo state, Brazil

Petrochirus diogenes, one of the largest hermit crabs in the western Atlantic, was studied in the north coast of the State of São Paulo, Brazil, to determine its breeding season through the analysis of gonad development at a macroscopic level. The analysis of 999 crabs collected from 1995 to 1999, based on four categories of gonad development, showed that males present gonads in the advanced stage during all seasons, and although females present developed ovaries all year as well, the peak incidence is in the summer. The peak of recruitment occurs in the winter.