Tomás A. Luppi

Distribution and habitat preferences of two grapsid crab species in Mar Chiquita Lagoon (Province of Buenos Aires, Argentina)

Cyrtograpsus angulatus andChasmagnathus granulata (Grapsidae) are the two dominant decapod crustacean species in the outer parts of Mar Chiquita Lagoon, the southernmost in a series of coastal lagoons that occur along the temperate Atlantic coasts of South America. Distribution and habitat preferences (water and sediment type) in these crab species were studied in late spring. There is evidence of ontogenetic changes in habitat selection of both species. Recruitment ofC. angulatus takes place mainly in crevices of tube-building polychaete (Ficopomatus enigmaticus) “reefs” and, to a lesser extent, also in other protected microhabitats (under stones). In the latter, mostly somewhat larger juveniles were found, suggesting that these are used as a refuge for growing individuals. Adults are most frequently found on unprotected muddy and sandy beaches.C. angulatus was found in all parts of Mar Chiquita Lagoon, including freshwater, brackish, and marine habitats.C. granulata, in contrast, was restricted to the lower parts of the lagoon, where brackish water predominates and freshwater or marine conditions occur only exceptionally. It showed highest population density on “dry mud” flats and inSpartina densiflora grassland, where it can build stable burrows and where high contents of organic matter occur in the sediment. Such habitats are characterized by mixed populations of juveniles (including newly settled recruits) and adults, males and females (including a high percentage of ovigerous). Unstable “wet mud” as well as stony sand were found to be inhabited by chiefly adult populations, with only few ovigerous females. In “dry mud” flats, the proportion of males increased vertically with increasing level in the intertidal zone, showing a significantly increasing trend also in their average body size. These observations may be explained by higher resistance of males, in particular of large individuals, to desiccation, salinity, and temperature stress occurring in the upper intertidal. However, an opposite, or no such, tendency was found in the distribution of ovigerous and non-ovigerous females, respectively. With increasing distance from the water edge, salinity increased and pH decreased significantly inC. granulata burrows, whereas temperature showed no consistent tendency within the intertidal gradient. A highly significant linear relationship (r=−0.794; P<0.001) between salinity and pH in water from crab burrows is described. This regression line is significantly different from one that had been observed in water from the lagoon, indicating consistently lower pH values at any salinity level in burrow water. This is interpreted as a result of crab and/or microbial respiration.

Hatching rhythms and dispersion of decapod crustacean larvae in a brackish coastal lagoon in Argentina

Mar Chiquita, a brackish coastal lagoon in central Argentina, is inhabited by dense populations of two intertidal grapsid crab species,Cyrtograpsus angulatus andChasmagnathus granulata. During a preliminary one-year study and a subsequent intensive sampling programme (November–December 1992), the physical properties and the occurrence of decapod crustacean larvae in the surface water of the lagoon were investigated. The lagoon is characterized by highly variable physical conditions, with oligohaline waters frequently predominating over extended periods. The adjacent coastal waters show a complex pattern of semidiurnal tides that often do not influence the lagoon, due to the existence of a sandbar across its entrance. Besides frequently occurring larvae (exclusively freshly hatched zoeae and a few megalopae) of the two dominating crab species, those of three other brachyurans (Plathyxanthus crenulatus, Uca uruguayensis, Pinnixa patagonica) and of one anomuran (the porcellanidPachycheles haigae) were also found occasionally. Caridean shrimp (Palaemonetes argentinus) larvae occurred in a moderate number of samples, with a maximum density of 800·m−3. The highest larval abundance was recorded inC. angulatus, with almost 8000°m−3. Significantly moreC. angulatus andC. granulata zoeae occurred at night than during daylight conditions, and more larvae (statistically significant only in the former species) during ebb (outflowing) than during flood (inflowing) tides. In consequence, most crab zoeae were observed during nocturnal ebb, the least with diurnal flood tides. Our data suggest that crab larvae do not develop in the lagoon, where the adult populations live, but exhibit an export strategy, probably based upon exogenously coordinated egg hatching rhythms. Zoeal development must take place in coastal marine waters, from where the megalopa eventually returns for settlement and metamorphosis in the lagoon. Significantly higher larval frequency ofC. granulata in low salinities (≤12‰) and at a particular sampling site may be related to local distribution patterns of the reproducing adult population. Unlike crab larvae, those of shrimp (P. argentinus) are retained inside the lagoon, where they develop from hatching through metamorphosis. They significantly prefer low salinity and occur at the lagoon surface more often at night. These patterns cannot be explained by larval release rhythms like those in brachyuran crabs, but may reflect diel vertical migrations to the bottom. It is concluded that osmotic stress as well as predation pressure exerted by visually directed predators (small species or life-cycle stages of estuarine fishes) may be the principal selection factors for the evolution of hatching and migration rhythms in decapod larvae, and that these are characteristics of export or retention mechanisms, respectively.

Effects of reduced salinities on development and bioenergetics of early larval shore crab, Carcinus maenas

The shore crab, Carcinus maenas L. (Portunidae), is a coastal and estuarine species, which can live and reproduce under brackish water conditions; freshly hatched larvae have been observed in the field at salinities below 15 parts per thousand. In the present laboratory study, the tolerance of hypo-osmotic stress was experimentally investigated in early larvae of a marine (North Sea) population of C. maenas reared at four different salinities (15, 20, 25, 32 parts per thousand). Two and 4 days after hatching, the Zoea I larvae were moult-staged microscopically, and their rates of respiration and growth (changes in dry weight, W, carbon, C, nitrogen, N, and hydrogen, H) were measured. Survival and development were monitored until the megalopa was reached: 15 parts per thousand did not allow for development beyond the first zoeal stage, while metamorphosis to the megalopa was reached at salinities greater than or equal to 20 parts per thousand. At 20 parts per thousand, development was significantly delayed and mortality enhanced as compared with 25 and 32 parts per thousand. Rates of growth and respiration decreased during exposure to reduced salinities less than or equal to 25 parts per thousand. Hence, the suppression of growth could not be explained as a consequence of enhanced metabolic losses per larva. Instead, a partial C budget indicates that the Zoea I larvae suffered from decreased capabilities of assimilating ingested and subsequently converting assimilated matter to tissue growth. Net growth efficiency (K-2, C-based) was at 25 and 32 parts per thousand initially high (> 60% during the postmoult and intermoult stages of the Zoea I moult cycle), but decreased during the later stages (down to less than or equal to 30% in premoult). An inverse pattern of C partitioning was observed at less than or equal to 20 parts per thousand, with initially low K-2 values (less than or equal to 21% during the first 2 days of the moult cycle), and a later increase (up to greater than or equal to 46% in premoult). Thus, larval growth was initially suppressed under conditions of reduced salinity, but this was later (during premoult) partially compensated for by an increase in C assimilation and K-2. Our observations indicate that Zoea I shore crab larvae react during the late stages of their moulting cycle less sensitively against reduced salinities than during postmoult and intermoult. This suggests that the transition between moult cycle stages C and D-0 may be a critical point for effects of hypo-osmotic stress, similarly as already known in relation to effects of nutritional stress. Negative effects were found also when freshly hatched Zoea I shore crab larvae were exposed only transitorily (for 24-72 h) to 20 parts per thousand, with significantly lower rates of survival, development, growth, respiration, and K-2. These effects increased with increasing duration of initial exposure to reduced salinity.

Experimental studies on predation and cannibalism of the settlers of Chasmagnathus granulata and Cyrtograpsus angulatus (Brachyura: Grapsidae)

The grapsid crabs Chasmagnathus granulata and Cyrtograpsus angulatus are considered as key species within the benthic communities of estuaries and brackish coastal lagoons in the southwestern Atlantic region. In controlled laboratory experiments, we studied the intensity of interspecific predation as well as intra- and intercohort cannibalism in setllers in relation to refuge availability, predator characteristics (species, size, sex, nutritional state), and the presence or absence of an alternative food source (Artemia nauplii). In both species, the intensity of intracohort cannibalism among recently settled crabs (instars I and II, “settlers”) was low, with ca. 5% mortality during 48 h experimental observation periods. Larger juveniles and adults of both species, by contrast, preyed heavily on the settlers. Predation was significantly reduced when refuges were available for the settlers. Hunger of the predators enhanced in general the predation rate. In the presence of alternative food, the consumption of settlers was significantly reduced. In C. granulata, adult females ate more settlers than the males, probably as a consequence of differences in the morphometric traits of their chelae. Cannibalism and predation by juvenile and adult crabs may play an important role in the regulation of recruitment success for both species and hence, in the structure of estuarine benthic communities.

Population structure of the South American Estuarine crab, Chasmagnathus granulatus (Brachyura: Varunidae) near the southern limit of its geographical distribution: comparison with northern populations

AbstractCrabs are among the most conspicuous and ecologically important invertebrates of the large intertidal zones that characterize estuarine and protected coastal areas in temperate regions. The habitat, population structure and breeding cycle of Chasmagnathus granulatus (Brachyura: Varunidae), a semiterrestrial burrowing crab endemic to the warm temperate coasts of the Southwestern Atlantic, were studied in San Antonio Bay (Argentina), near the southern limit of its range. San Antonio Bay has no freshwater input, winter is relatively colder, and summer warmer, than northern habitats of this species. Crabs lived both in vegetated and unvegetated zones, but density and sex ratio varied among dates and zones. The maximum observed density was 136 crabs/m2 , the maximum carapace width (CW) was 32 mm (males) and 29.8 mm (females), ovigerous females were found only in November and January, and the smallest ovigerous female measured 17 mm CW. The population structure, spatial distribution, and recruitment pattern of C. granulatusdid not differ between San Antonio Bay and northern habitats. The higher density, smaller maximum size and shorter reproductive cycle observed in San Antonio cannot be atributed to changes associated with a latitudinal cline and other factors, such as thermal amplitude and food availability, need to be studied.

Population Structure of the SW Atlantic Estuarine Crab Neohelice granulata Throughout Its Range: a Genetic and Morphometric Study

Phylogeographic patterns of the SW Atlantic estuarine crab Neohelice granulata were examined using mitochondrial DNA cytochrome oxidase I sequences and analyzed together with morphometric data. Specimens were sampled during a 4-year period (2001–2004) from 11 localities encompassing the full distributional range of this species along the SW Atlantic (22°57′ S to 42°25′ S). DNA sequences were obtained from 69 individuals belonging to seven localities, and morphometric variation in 12 continuous characters was analyzed for 646 crabs from ten localities. Strong genetic differentiation, consistent with a pattern of isolation-by-distance, was detected among all localities indicating that gene flow occurs mainly between neighboring populations. Analyses of molecular variance showed genetic subdivision between the southern (Argentina) and the northern (Brazil) sites, suggesting restricted gene flow at a regional scale. The genetic structure of this species could be divided into two distinct groups due to a limited gene flow between southern and northern regions as a consequence of larval dispersal patterns. Coastal currents in the vicinity of the Rio de la Plata likely act as a barrier to dispersal within the species range. Moreover, genetic data indicate that populations of N. granulata might have undergone a northward demographic expansion since the late Pleistocene. The morphometric analysis showed no geographical pattern of morphological differentiation, although there were differences among sampling sites.

Receptivity of female Neohelice granulata (Brachyura, Varunidae): different strategies to maximize their reproductive success in contrasting habitats

The extent of the receptive period may determine the mating strategies employed by female crabs to obtain mates. Here, we studied the receptivity of female Neohelice granulata (Dana, 1851) in the laboratory, including the form of the vulvae and the anatomy of the seminal receptacle (SR). We examined the factors that influence the duration of receptivity by comparing two populations inhabiting contrasting habitats: Mar Chiquita Coastal lagoon (MCL), which is an oligo-polyhaline estuary, and San Antonio Oeste (SAO), which is an eu-hyperhaline marine bay. Non-receptive females have immobile vulva opercula, while receptive females have mobile opercula. Histological sections of the SR showed that the degree of epithelium secretions was associated with the receptive stage of females, and they may be involved in the maintenance of viable sperm and in the dehiscence of spermatophores. The existence of a special tissue at the junction of the oviduct and the SR was described and proposed as an internal mechanism influencing the timing of ovulation. The duration of receptivity was dependent on the SR load and the capacity to lay eggs. Thus, females with empty SR exhibited longer receptivity and did not lay eggs, while those with full SR exhibited shorter receptivity and always laid eggs. Interpopulation differences showed that females from SAO had shorter receptivity and heavier SR and laid eggs more frequently than females from MCL. Based on our results, we suggest that N. granulata females can adjust the duration of their receptivity and control the moment of fertilization according to different internal mechanisms related to the morphology of the vulvae, the fullness of the SR and anatomical attributes of the SR. An important consequence of this control is greater sperm competition. The extent of the receptive period and the number of times that a female could become receptive in a single reproductive season may also depend on the habitat characteristics.

Stage-dependent interactions between intertidal crabs: from facilitation to predation

Large parts of the south-western Atlantic soft bottom intertidals are inhabited by the burrowing crab Neohelice granulata (previously known as Chasmagnathus granulatus ) and the mud crab Cyrtograpsus angulatus , but adults rarely coexist in the same microhabitat. We describe the influence of burrows of N. granulata on the recruitment dynamic of C. angulatus and the effects of different ontogenetic stages of N. granulata on survival of recruits of C. angulatus . A two summer sampling shows that N. granulata burrows facilitate settlement of both species. To evaluate the mortality of recruits inside burrows, we performed a field experiment with juvenile and adult crab exclusion cages and inclusion of juveniles or adults of N. granulata . The results showed differences in crab sizes between treatments, due to predatory interactions that depended on prey size. When only juveniles of N. granulata were present, the higher mortality of C. angulatus was observed in smaller crabs. However when adults of N. granulata were present, larger juvenile crabs are the ones that suffered the highest mortality. These results show that adults are preying upon larger juveniles (of both species) reducing the mortality of the smaller ones that are preyed by the larger ones. We also measured emigration from these burrowing assemblages using bidirectional pitfall traps which showed that C. angulatus juveniles are leaving the burrows towards the subtidal. Here we demonstrate that N. granulata have positive and negative effects on the settlement of C. angulatus . These results highlight the importance of including stage-dependence relationships to analyse species interaction in marine ecology field works.

LARVAL MORPHOLOGY OF THE SOUTHWESTERN ATLANTIC MUD CRAB PANOPEUS MERIDIONALIS (DECAPODA: BRACHYURA: PANOPEIDAE) DESCRIBED FROM LABORATORY-REARED MATERIAL

Abstract The panopeid mud crab Panopeus meridionalis is known to inhabit the intertidal zone of the estuarine temperate waters of the southwestern Atlantic, from Montevideo, Uruguay, to Mar del Plata, Argentina. The complete larval development of this species is described from laboratory-reared material and compared with known larvae of Panopeus.

Epibiosis on eggs and brooding care in the burrowing crab Chasmagnathus granulatus (Brachyura:Varunidae): comparison between mudflats and salt marshes

Chasmagnathus granulatus is a semiterrestrial intertidal burrowing crab that inhabits both the unvegetated mudflats and the cordgrass ( Spartina densiflora ) salt marshes in Mar Chiquita Lagoon (Argentina), where it is considered the ecologically key species. The mass of C. granulatus eggs incubated by females is colonized by epibiotic micro-organisms and accumulates detritus. The type of epibionts that use eggs as a substrate, the infestation degree, the maternal care behaviour and the protection of the incubation chamber were compared between females living on mudflats and on Spartina -dominated areas. In both places, the epibiosis by bacteria and filamentous fungi and peritrichid colonial ciliate was significantly higher in the periphery than in the centre of the brood mass. The accumulation of detritus was higher in the periphery in mudflat females but not in salt marsh females. Moreover, the level of detritus was significantly higher in mudflat than in salt marsh females only in the periphery of the brood. The infestation level of bacteria and fungi, and peritrichids, increased throughout the embryonic development only in mudflat females. The periphery of the brood mass was significantly more contaminated in mudflat than in marsh females, while the central region of the brood mass did not differ between habitats. The pleopods were significantly more contaminated by bacteria and filamentous fungi and peritrichid colonial ciliates in premoult females than in postmoult females, independently from the collection site. The percentage of females with abnormal embryos was significantly higher in mudflats (26.7%) than in marshes (12.3%). Females with late embryos spent more time flapping the abdomen and probing the embryos with the chela. Non-ovigerous females did not perform specific maternal care activities. The volume of brood mass both in early or late stage of development is greater than that of the incubation chamber and, consequently, peripheral embryos are more exposed.