Luis Giménez

Relationships among salinity, egg size, embryonic development, and larval biomass in the estuarine crab Chasmagnathus granulata Dana, 1851.

We studied interrelationships between initial egg size and biomass, duration of embryogenesis at different salinities, and initial larval biomass in an estuarine crab, Chasmagnathus granulata. Ovigerous females were maintained at three different salinities (15 per thousand, 20 per thousand and 32 per thousand); initial egg size (mean diameter), biomass (dry weight, carbon and nitrogen) as well as changes in egg size, embryonic development duration, and initial larval biomass were measured.Initial egg size varied significantly among broods from different females maintained under identical environmental conditions. Eggs from females maintained at 15 per thousand had on average higher biomass and larger diameter. We hypothesise that this is a plastic response to salinity, which may have an adaptive value, i.e. it may increase the survivorship during postembryonic development. The degree of change in egg diameter during the embryonic development depended on salinity: eggs in a late developmental stage were at 15 per thousand significantly larger and had smaller increment than those incubated at higher salinities. Development duration was longer at 15 per thousand, but this was significant only for the intermediate embryonic stages. Initial larval biomass depended on initial egg size and on biomass loss during embryogenesis. Larvae with high initial biomass originated either from those eggs that had, already from egg laying, a high initial biomass (reflecting individual variability under identical conditions), or from those developing at a high salinity (32 per thousand), where embryonic biomass losses were generally minimum. Our results show that both individual variability in the provisioning of eggs with yolk and the salinity prevailing during the embryonic development are important factors causing variability in the initial larval biomass of C. granulata, and thus, in early larval survival and growth.

Small changes in pH have direct effects on marine bacterial community composition: a microcosm approach

As the atmospheric CO2 concentration rises, more CO2 will dissolve in the oceans, leading to a reduction in pH. Effects of ocean acidification on bacterial communities have mainly been studied in biologically complex systems, in which indirect effects, mediated through food web interactions, come into play. These approaches come close to nature but suffer from low replication and neglect seasonality. To comprehensively investigate direct pH effects, we conducted highly-replicated laboratory acidification experiments with the natural bacterial community from Helgoland Roads (North Sea). Seasonal variability was accounted for by repeating the experiment four times (spring, summer, autumn, winter). Three dilution approaches were used to select for different ecological strategies, i.e. fast-growing or low-nutrient adapted bacteria. The pH levels investigated were in situ seawater pH (8.15–8.22), pH 7.82 and pH 7.67, representing the present-day situation and two acidification scenarios projected for the North Sea for the year 2100. In all seasons, both automated ribosomal intergenic spacer analysis and 16S ribosomal amplicon pyrosequencing revealed pH-dependent community shifts for two of the dilution approaches. Bacteria susceptible to changes in pH were different members of Gammaproteobacteria, Flavobacteriaceae, Rhodobacteraceae, Campylobacteraceae and further less abundant groups. Their specific response to reduced pH was often context-dependent. Bacterial abundance was not influenced by pH. Our findings suggest that already moderate changes in pH have the potential to cause compositional shifts, depending on the community assembly and environmental factors. By identifying pH-susceptible groups, this study provides insights for more directed, in-depth community analyses in large-scale and long-term experiments.

Phenotypic links in complex life cycles: conclusions from studies with decapod crustaceans

I review studies on decapod crustaceans to draw conclusions about the importance of effects of past environmental conditions on development, phenotype, performance, and survival in animals. I consider 3 critical points of the life cycle: the allocation of reserves into eggs, the hatching of larvae, and metamorphosis from the larval to the juvenile phase. Biomass allocated to eggs varies among females as a response to changes in environmental conditions. These variations are propagated to the larval stages, influencing the biomass at hatching, subsequent larval developmental pathways, and survival during periods of limited starvation. Suboptimal conditions experienced by embryos increase the loss of mass during embryogenesis; size or biomass of the juvenile is either positively or negatively correlated with initial biomass. Positive correlations may be the normal pattern; negative correlations occur when individuals hatched with low initial biomass follow developmental pathways that lead to increased biomass at metamorphosis. In estuarine crabs, salinity experienced by embryos leads to salinity acclimation in early larval stages. Phenotypic links originate as transgenerational effects that propagate to the juvenile stages. There are least 3 types of effects: disruption of physiological processes; direct adaptive responses; and indirect consequences of adaptive mechanisms. All types appear within a species; they are produced as a response to a single environmental factor. Variability in phenotype remains latent and is expressed in terms of survival according to the environmental conditions experienced by a particular stage. The fate of individuals is thus affected by interactions between their immediate developmental processes and their environmental history.

Effects of prehatching salinity and initial larval biomass on survival and duration of development in the zoea 1 of the estuarine crab, Chasmagnathus granulata, under nutritional stress

Abstract The effects of individual larval biomass, and salinity experienced during embryogenesis (i.e., prehatching salinity) on starvation tolerance and growth of zoea 1 of the estuarine crab (Chasmagnathus granulata) were evaluated in laboratory experiments. Freshly hatched zoeae 1 were obtained from broods maintained at three salinities (15‰, 20‰ and 32‰), and cultured at 20‰ under different initial feeding periods and subsequent food deprivation (“point of reserve saturation” experiment: PRS) or under initial periods of food deprivation and subsequent feeding (point of no return experiment: PNR). Another group of larvae were used for determination of biomass (dry weight, carbon, and nitrogen) of zoea 1. Larval survival and duration of development depended on the length of feeding period: no larvae reached the second instar under complete starvation; survival was higher and duration of development shorter as the feeding period lengthened. After different initial feeding periods (PRS experiment), zoeae 1 that hatched from eggs incubated at the prehatching salinities of 15‰ and 20‰ showed higher survival and shorter duration of development than those at 32‰. Prehatching salinity also affected the amount of reserves accumulated during the first 2 days after hatching, with larvae from 15‰ and 20‰ showing the highest percentage of total accumulation of carbon and nitrogen. Initial larval biomass did not affect survival, but it had a slight effect on duration of development, with larger larvae (in terms of biomass) developing faster. After different initial starvation periods (PNR experiment), prehatching salinity did not affect survival, but it affected duration of development: larvae from 15‰ and 20‰ reached the second instar earlier. Variability in survival and duration of development was explained in part by among-brood variability in initial larval biomass: larvae with higher biomass showed higher survival and shorter duration of development. Thus, C. granulata, survival and duration of development under food stress depend on the interaction between environmental conditions experienced before and after hatching (pre- and posthatching factors, respectively).

Relationships between habitat conditions, larval traits, and juvenile performance in a marine invertebrate

Environmental effects on sets of traits of organisms can have important consequences on populations. In marine species with complex life cycles, plastic responses in larval traits can affect size at metamorphosis and juvenile growth. These traits can ultimately affect survival and recruitment. Here, I evaluate links between pelagic conditions, natural variations in larval physiological traits (duration of development, size of the megalopa stage), size at metamorphosis, and juvenile growth in the shore crab Carcinus maenas. I sampled settling larvae (megalopae) of C. maenas, daily, during the settlement season of three consecutive years, in the island of Helgoland (North Sea, German Bight). Megalopae were brought to the laboratory where postmetamorphic growth was evaluated under different larval food environments. Traits varied considerably at several temporal scales; variation was not consistent among years. However, the size at metamorphosis was always larger at the start of the settlement season. Some variability correlated with predicted changes in transport processes and dynamics of plume fronts around the study area. Positive relationships between predicted periods of formation of plume fronts, settlement rates, and size at metamorphosis suggest a link between oceanographic conditions and individual traits. Size at metamorphosis explained juvenile growth rates: during the first five juvenile stages, larger individuals grew faster and reached larger size at stage (that is, at any stage observed). Megalopae experiencing ad libitum food conditions grew faster as juveniles than those deprived of food, showing that size at metamorphosis and juvenile growth respond plastically to variations in the larval food environment. I conclude that juvenile growth rate in C. maenas can be affected by natural environmental conditions experienced by larval stages; this effect is mediated by changes in the size at metamorphosis. Most likely the most important factor operating on the size at metamorphosis was food (availability/quality). Since juveniles of this species attain a refuge in size from cannibals, changes in size at metamorphosis could codetermine the chances of postsettlement survival. Further consideration of physiological traits and their consequences at the level of populations should lead to a better understanding of how pre-and postmetamorphic processes interact and explain variability in recruitment in species with complex life cycles.

Scale-dependent patterns of macrofaunal distribution in soft-sediment intertidal habitats along a large-scale estuarine gradient

We investigated the pattern of distribution of intertidal soft-bottom fauna in streams and lagoons of the Uruguayan coast at three spatial scales. The Río de la Plata and the Atlantic Ocean produce on this coast a large-scale gradient in salinity, defining a freshwater (west), an estuarine (central) and a marine (east) region. Within each region, there are several streams and coastal lagoons (sites) that define a second scale of variability. A third scale is given by intertidal gradients within each site. Species richness and total abundance was low in the freshwater west region and high in the central and east regions. The community in the west region was characterized by the clam Curbicula fluminea; in the other regions, it was dominated mainly by the polychaete Heteromastus similis. The polychaete Nephtys fluviatilis was more abundant in the east region, while another polychaete, Laeonereis acuta, characterized the central region. Sediment fractions did not vary significantly at this scale. At the scale of the sites, species richness and total macrofaunal abundance were higher in coastal lagoons than in streams. Coarse sands were more common in coastal lagoons, while medium and fine sand characterized the sediment in streams. Within each site, species richness and total abundance increased towards the lower intertidal level; the macrofauna of the upper levels were a subsample of the fauna occurring at the lower levels. There was also a significantly lower proportion of fine sand at the upper level. At regional scales, the observed patterns may be indirectly or directly related to the gradient in salinity, through differential physiological tolerance to osmotic stress. At the scale of the sites, variability may be explained mainly by geomorphological and sedimentological differences between lagoons and streams. Variation among levels may be related to gradients in desiccation, colonization and predation.

Effects of reduced salinity on the biochemical composition (lipid, protein) of zoea 1 decapod crustacean larvae

Effects of reduced salinities on dry weight (DW) and biochemical composition (total lipid and protein contents) of zoea 1 larvae were evaluated in four decapod crustacean species differing in salinity tolerance (Cancer pagurus, Homarus gammarus, Carcinus maenas, Chasmagnathus granulata). The larvae were exposed to two different reduced salinities (15xand 25xin C. granulata ,2 0x and 25x in the other species) for a long (ca. 50% of the zoea 1 moulting cycle) or a short period (16 h, starting at ca. 40% of the moulting cycle), while a control group was continually maintained in seawater (32x). In general, the increments in dry weight, lipid and protein content were lower at the reduced salinities than in the control groups. In the zoea 1 of H. gammarus (stenohaline) and C. pagurus (most probably also stenohaline), the lipid and protein contents varied greatly among treatments: larvae exposed to low salinities exhibited very low lipid and protein contents at the end of the experiments compared to the controls. In some cases, there were negative growth increments, i.e. the larvae had, after the experimental exposure, lower lipid and protein contents than at the beginning of the experiment. C. maenas (moderately euryhaline) showed a lower variation in protein and lipid content than the above species. The zoea 1 of C. granulata (fairly euryhaline) showed the lowest variability in dry weight, protein and lipid content. Since salinity tolerance (eury- v. stenohalinity) is associated with the osmoregulatory capacity, our results suggest a relationship between the capability for osmoregulation and the degree of change in the biochemical composition of larvae exposed to variable salinities.

Potential effects of physiological plastic responses to salinity on population networks of the estuarine crab Chasmagnathus granulata

Abstract.Chasmagnathus granulata is a South American crab occurring in estuarine salt marshes of the Brazilian, Uruguayan and Argentine coasts. Life history is characterized by an export strategy of its larval stages. I reviewed information on experimental manipulation of salinity during embryonic and larval development (pre- and posthatching salinities), and on habitat characteristics of C. granulata in order to determine potential effects of larval response to salinity in the field and to suggest consequences for the population structure. Local populations are spread over coastal areas with different physical characteristics. Benthic phases occupy estuaries characterized by different patterns of salinity variation, and release larvae to coastal waters characterized by strong salinity gradients. The zoea 1 of C. granulata showed a strong acclimatory response to low salinity. This response operated only during the first weeks of development (during zoeae 1 and 2) since subsequent larval survival at low posthatching salinities was consistently low. Larvae developing at low salinity frequently followed a developmental pathway with five instead of four zoeal stages. The ability to acclimate and the variability in larval development (i.e. the existence of alternative developmental pathways) could be interpreted as a strategy to buffer environmental variability at spatial scales of local or population networks. Early survivorship and production of larvae may be relatively high across a rather wide range of variability in salinity (5–32‰). Plastic responses to low salinity would therefore contribute to maintain a certain degree of population connectivity and persistence regardless of habitat heterogeneity.

Swimming ability and burrowing time of two cirolanid isopods from different levels of exposed sandy beaches

Abstract Most of the macroinfauna from sandy beaches is highly mobile, emerging out of the sediment when the tide rises, and using the swash to migrate up and down the beach face or feed (searching for prey or carrion). After swash excursions, they usually burrow back into the sediment, maintaining zonation at low tide. Therefore, the different species abilities to emerge, move around and burrow under different swash climates and sediment conditions are expected to influence observed distribution patterns. Nonetheless, few attempts have been made to understand behavioral mechanisms of these organisms in moving fluids. In this study, we used a flume tunnel to investigate the orientation, swimming ability and burrowing time of two similar species of cirolanid isopods, Excirolana armata Dana and Excirolana braziliensis Richardson, under current velocities ranging from 5 to 30 cm·s −1 . E. armata inhabits middle levels of dissipative to intermediate beaches, while E. braziliensis is found towards the upper level of a wider range of beach states. Both species oriented countercurrent above a threshold velocity, which turned out to be significantly lower for E. armata than for E. braziliensis . E. armata proved to be a stronger swimmer as shown by the higher velocities surmounted, and the less drags experienced at the highest current velocity. E. armata also burrowed faster than E. braziliensis . Burrowing time was affected by sediment grain size and water content, but not by water flow. Once organisms managed to begin burrowing under different flow conditions, they were not affected by current velocity. Nonsaturated sand precluded burial, while coarse sand retarded it. Differences in the observed patterns of across-beach distribution may thus be the result of species-specific behavioral responses to swash climate, manifested in swimming ability, burying and orientation in directional flows.

Fish assemblage in a temperate estuary on the uruguayan coast: seasonal variation and environmental influence

The seasonal dynamics of the fish community in the Pando estuary on the Uruguayan coast were studied in relation to environmental sampled monthly between May 2002 and June 2003. Individuals collected were identified, and classified into stages (juveniles, adults) and functional groups. Relationships between community dynamics and environmental variables were evaluated using uni- and multivariate techniques. Twenty-one species, mostly freshwater stragglers, estuarine and marine migrants were collected. The most abundant species were Micropogonias furnieri, Mugil platanus, Paralichthys orbignyanus and Brevoortia aurea and were represented by juveniles. The community varied seasonally with rapid shifts in spring and autumn associated with changes in temperature and salinity. Significant correlations between abundance and temperature may be related to the timing of life cycle events. In this estuary, the salinity appears to play a key role in the functional structure and in the use of the habitat by juveniles. This is relevant for the definition of estuaries as nursery areas: this definition is context-dependent and is determined by the salinity conditions.