Short-term response of deposit-feeders to an increase of the nutritive value of the sediment through seasons in an intertidal mudflat (Western Mediterranean, Italy)

In coastal intertidal flats, assemblages of deposit-feeders often vary seasonally and according to the availability of food in sediments. In the mudflat where this study has been done, the deposit-feeder assemblage changed after a sharp increase of fresh detritus, which occurred in summer 1997. The number of the animals increased and abundances of some worms (Streblospio shrubsolii, Desdemona ornata and Capitella cf. capitata) were correlated with the nutritive value of the sediment. In addition, there was evidence that increasing food in summer would increase recruitment of juveniles and adults of S. shrubsolii.In this paper, the concentration of food was manipulated in situ in sediments of an intertidal mudflat in different seasons. Food of high nutritive value was added to small plots of sediments to test hypotheses about effects on the distribution of deposit-feeders. It was predicted that addition of food would change composition and abundance of the assemblage and densities of the deposit-feeders that were correlated to the availability of food. Furthermore, it was tested whether juveniles and adults of S. shrubsolii were affected by the availability of food consistently through seasons.Densities of deposit-feeders increased in plots where the sediment was organically enriched. The composition of the assemblage, however, was not affected. S. shrubsolii, D. ornata and C. cf. capitata responded to the addition of food. However, the nutritive value of the sediment affected the dynamics of these animals variably through seasons. Abundances increased in the experimental treatment E in winter and spring for D. ornata and in winter and autumn for C. cf. capitata. Furthermore, in summer and autumn juveniles and adults of S. shrubsolii were more abundant in plots where food was added.     

Predation on 0-group and older year classes of the bivalve Macoma balthica: interaction of size selection and intertidal distribution of epibenthic predators

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea.Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma.Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation.     

The effect of wrack composition and diversity on macrofaunal assemblages in intertidal marine sediments

Wrack (dead, washed-up seaweed and seagrass) buried in soft substrata may increase the organic content and alter the physical structure of sediments. These effects may influence the composition and structure of macrofaunal assemblages in the sediment. Such influences can be expected to vary according to the type and amount of wrack as well as the presence of invasive seaweeds in the wrack. In this study, we deliberately buried different amounts of the invasive species Sargassum muticum in isolation or mixed to the native species Ulva sp. and Fucus vesiculosus, in two intertidal sandflats to test some hypotheses about the response of macrofaunal assemblages. We tested whether (1) diversity of detritus (i.e. different mixtures), and (2) the amount of detritus of S. muticum influenced the composition and the relative abundance of macrofaunal assemblages. We also assessed whether the sediment organic carbon and the biomass of benthic microalgae varied depending on the diversity of detritus and the amount of detritus of S. muticum. Finally, we tested if these effects of wrack were consistent across sites. Results indicated that buried wrack affected the composition and structure of macrofaunal assemblages in short-term (i.e. 4 weeks), but there were no differences depending on detritus diversity or the amount of S. muticum. In addition, sediment organic matter and microalgal biomass were not affected by the addition of wrack. They instead varied greatly among small spatial scales (i.e. plots). Wrack composition or abundance of the invasive species S. muticum played thus a small role in shaping the structure of macrofaunal assemblages or the biomass of benthic microalgae in these intertidal sediments, probably because these sediments are frequently affected by various inputs of organic matter and benthic assemblages are already adapted to organically enriched sediments.     

Algae, macrofaunal assemblages and temperature: a quantitative approach to intertidal ecosystems of Iceland

Algae and the associated macrofauna in two Icelandic intertidal ecosystems under cold and warm influence, respectively, were studied with respect to algae-macrofauna relationships and a possible effect of temperature on community structure. Two sites in Iceland were selected, Sandgerdi ligthhouse (64°8′N 22°40′W) on the southwestern coast, and Grimsey Island (66°33′N 18°04′W), in the north, on the Arctic Circle, where sea temperature is considerably lower (5° approximately). The biomass of algae and the number of species of algae and macrofauna were higher in Sandgerdi than in Grimsey, and the patterns of diversity, evenness, biomass and abundance also differed between the sites. In the intertidal zone of Sandgerdi, a total of 28 species of algae and 45 species of macrofauna were identified whereas only 16 algal species and 27 macrofaunal species were found in Grimsey. Canonical correspondence analysis (CCA) using algal biomass as the environmental variable were conducted, and revealed significant relationships between algae composition and the associated macrofauna; some macrofauna taxa showed specific trophic or refuge relationships with algal species. According to the CCA, Corallina officinalis showed the highest correlation with macrofaunal assemblages in both study sites. However, correlations between macrofauna and other algae differed between Grimsey and Sandgerdi. The present study, together with additional observations in Greenland waters, shows a general decrease of species richness and diversity towards the north which may primarily be due to the temperature regime.     

Occurrence of epibenthic dinoflagellates in relation to biotic substrates and to environmental factors in Southern Mediterranean (Bizerte Bay and Lagoon,Tunisia): An emphasis on the harmful Ostreopsis spp., Prorocentrum lima and Coolia monotis

Harmful events associated with epibenthic dinoflagellates, have been reported more frequently over the last decades. Occurrence of potentially toxic benthic dinoflagellates, on the leaves of two magnoliophytes (Cymodocea nodosa and Zostera noltei) and thalli of the macroalgae (Ulva rigida), was monitored over one year (From May 2015 to April 2016) in the Bizerte Bay and Lagoon (North of Tunisia, Southern Mediterranean Sea). The investigated lagoon is known to be highly anthropized. This is the first report on the seasonal distribution of epibenthic dinoflagellates hosted by natural substrates, from two contrasted, adjacent coastal Mediterranean ecosystems. The environmental factors promoting the development of the harmful epibenthic dinoflagellates Ostreopsis spp., Prorocentrum lima and Coolia monotis were investigated. The highest cell densities were reached by Ostreopsis spp. (1.9 × 10³ cells g⁻¹ FW, in October 2015), P. lima (1.6 × 10³ cells g⁻¹ FW, in June 2015) and C. monotis (1.1 × 10³ cells g⁻¹ FW, in May 2015). C. nodosa and Z. noltei were the most favorable host macrophytes for C. monotis (in station L2) and Ostreopsis spp. (in station L3), respectively. Positive correlations were recorded between Ostreopsis spp. and temperature. Densities of the epibenthic dinoflagellates varied according to the collection site, and a great disparity was observed between the Bay and the Lagoon. Maximum concentrations were recorded on C. nodosa leaves from the Bizerte Bay, while low epiphytic cell abundances were associated with macrophytes sampled from the Bizerte Lagoon. The observed differences in dinoflagellate abundances between the two ecosystems (Bay-Lagoon) seemed not related to the nutrients, but rather to the poor environmental conditions in the lagoon.     

Influence of sediment on burrowing by the soldier crab Mictyris longicarpus Latreille

This study analyses the spatial distribution of burrowing by the soldier crab Mictyris longicarpus (Mictyridae) in relation to the topography of intertidal sandflats. Four different locations were sampled in Port Jackson and Botany Bay (Sydney, New South Wales, Australia). In each location, abundances of crabs and the proportion of juvenile crabs varied among sites 50 m² and tens of metres apart, but not between high- and lowshore levels, nor among shores. Topographic formations, referred to as humps and depressions, were common on the flats and were of the same order of size and spacing as the sites, which showed variation in numbers/sizes of crabs. Manipulative experiments were done in one of the locations to distinguish between the hypotheses that the spatial distribution of the crabs varied according to (1) topography, or (2) the composition of sediment in the different topographic features. In the first experiment, small plots (0.09 m² and 20 cm deep) of sediment were transplanted between humps and depressions, including appropriate controls for disturbance and translocation. The numbers and sizes of crabs colonizing the experimental plots were then measured. Differences in abundance among plots were found higher on the shore. Therefore, the experiment was repeated at highshore levels with increased replication. There were more crabs in depressions than in humps. The proportion of juveniles varied according to the type of sediment, but only in depressions. Heterogeneity and sediment were, therefore, important for explaining some of the spatial variability of the abundances of the crabs.     

The photosynthetic performance of the tropical seagrass Halophila ovalis in the upper intertidal

Some ecophysiological adaptation strategies of the tropical seagrass Halophila ovalis were investigated with respect to this plant's ability to grow in the upper intertidal in either monospecific pools (but not together with other intertidal species) or emergent and exposed to high temperatures and irradiances during several hours every day. It was found that Halophila ovalis could raise the pH in simulated pools to 8.6, while the two other major (biomass wise) intertidal seagrasses raised the pH to 8.8 (Cymodocea rotundata) and 9.2 (Thalassia hemprichii). In situ, midday pH values of 8.5, 8.7 and 9.0 were recorded in pools inhabiting the three species, respectively. It was further found that photosynthetic electron transport rates (ETR) measured continuously in situ for one leaf during a diurnal cycle followed the irradiance up to a daily maximal value of 2200 μmol photons m⁻² s⁻¹, but dropped by ca. 50% when the leaf became air exposed just before noon. This drop in ETR upon emergence was verified by “point measurements”, while ETRs of leaves that grew in small pools formed as the tide receded followed the irradiance more closely. Similarly, F_v/F_m measured after 15 min of dark adaptation decreased by ca. 50% in emergent leaves during midday, but maintained higher daily values in the submerged leaves.     

Evidences of habitat displacement between two common soft-bottom SW Atlantic intertidal crabs

The intertidal burrowing crab Chasmagnathus granulatus Dana is the dominant species in soft sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 41°S) where it produces dense and extensive burrowing beds. The mud crab Cyrtograpsus angulatus Dana coexists with Ch. granulatus in this area, but it also inhabits areas to the south (northern and central Argentinean Patagonia). A survey covering both areas showed that C. angulatus rarely live in burrows when coexisting with Ch. granulatus, but form large burrowing beds when not coexisting with Ch. granulatus. When both species coexisted, burrowing beds of C. angulatus are restricted to sandy-muddy areas. Only rarely are burrows of C. angulatus found within Ch. granulatus beds. However, when Ch. granulatus were experimentally excluded within their burrowing beds, new settlers of C. angulatus made burrows and maintained them until they reached large size. Paired (inside and outside Ch. granulatus burrowing bed) sampling during high tide using beach nets showed that C. angulatus rarely venture inside the Ch. granulatus crab beds. Other field experiments showed that adults Ch. granulatus always displace C. angulatus from burrows. Furthermore, in several sites located south of the limit of distribution of Ch. granulatus at the Patagonian coast, soft bare intertidals are dominated by burrowing beds of C. angulatus mixed with the congener C. altimanus Dana. Together, these evidences suggest that the mud crab C. angulatus is displaced from soft bottom areas by the burrowing crab Ch. granulatus. It is an example of competitive exclusion through aggressive interference in soft-bottom habitats when the shared resource is the access to sediment surface, a two-dimensional well-defined resource.     

Response of predatory snails to a novel prey following the geographic range expansion of an intertidal barnacle

During a geographic range expansion, the success of a species colonizing a new region may be influenced by how resident predators respond to this novel prey. The volcano barnacle Tetraclita rubescens has recently expanded its range > 300 km northward along the coast of California, USA. In the historic portion of its range, Tetraclita is preyed upon by a southern dogwhelk, Nucella emarginata. In newly colonized northern habitats, Tetraclita has joined an assemblage of mid-intertidal barnacles preyed upon by two species of northern dogwhelks (N. ostrina and N. canaliculata). Using a combination of field enclosures and laboratory experiments, we tested the impacts of northern whelks on volcano barnacles relative to other barnacle species. In the field, juvenile Tetraclita were rarely consumed. In laboratory trials with larger barnacles, Tetraclita were nearly immune to drilling by northern whelks, perhaps due to superior morphological defenses. In all experiments, prior encounters with Tetraclita did not appear to increase the frequency of predation on this barnacle; northern whelk species rarely preyed on Tetraclita even when collected from a site where this barnacle was abundant. In contrast, the southern whelk N. emarginata readily preyed on Tetraclita in laboratory trials, perhaps reflecting a longer shared evolutionary history between these species. Overall, our results suggest that the success of a population established beyond a species' historical range boundary may be influenced by the functional traits of that species in relation to the composition and evolutionary history of the surrounding community.     

In and out: an analysis of epibiotic vs periplasmic bacterial predators

Bdellovibrio and like organisms (BALO) are obligate predators of Gram-negative bacteria, belonging to the α- and δ-proteobacteria. BALO prey using either a periplasmic or an epibiotic predatory strategy, but the genetic background underlying these phenotypes is not known. Here we compare the epibiotic Bdellovibrio exovorus and Micavibrio aeruginosavorus to the periplasmic B. bacteriovorus and Bacteriovorax marinus. Electron microscopy showed that M. aeruginosavorus, but not B. exovorus, can attach to prey cells in a non-polar manner through its longitudinal side. Both these predators were resistant to a surprisingly high number of antibiotic compounds, possibly via 26 and 19 antibiotic-resistance genes, respectively, most of them encoding efflux pumps. Comparative genomic analysis of all the BALOs revealed that epibiotic predators have a much smaller genome (ca. 2.5 Mbp) than the periplasmic predators (ca. 3.5 Mbp). Additionally, periplasmic predators have, on average, 888 more proteins, at least 60% more peptidases, and one more rRNA operon. Fifteen and 219 protein families were specific to the epibiotic and the periplasmic predators, respectively, the latter clearly forming the core of the periplasmic ‘predatome'', which is upregulated during the growth phase. Metabolic deficiencies of epibiotic genomes include the synthesis of inosine, riboflavin, vitamin B6 and the siderophore aerobactin. The phylogeny of the epibiotic predators suggests that they evolved by convergent evolution, with M. aeruginosavorus originating from a non-predatory ancestor while B. exovorus evolved from periplasmic predators by gene loss.     

Regeneration of lost siphon tissues in the tellinacean bivalve Nuttallia olivacea

The inhalant siphon of the tellinacean bivalve Nuttallia olivacea is an important prey item for juvenile stone flounder Platichthys bicoloratus in estuaries in Japan. We examined quantitative siphon regeneration of N. olivacea in rearing experiments of siphon-removed bivalves (> 30 mm shell length) both in the laboratory and in their natural habitat. Under laboratory conditions, siphon-removed bivalves regenerated lost tissues quantitatively at 15 and 25 °C 1 mo after siphon removal, although regeneration was incomplete. A 3-mo caging experiment in the field showed that great regeneration occurred in siphon-removed bivalves. However, the siphon weight of removed bivalves was significantly smaller than that of non-amputated bivalves, suggesting the incomplete regeneration. In a 1-mo caging experiment, bivalves that had approximately 15% of their siphons amputated were selected at some intervals to illustrate the quantitative regeneration process. Estimated daily siphon production was remarkably high only a few days after amputation. It decreased greatly thereafter, but regeneration was not completed within 30 d. These results indicate that bivalves regenerate siphons rapidly just after losing siphon tissues and then regeneration is slowed down before it is completed.     

Environmental and behavioural factors affecting activity in the intertidal gastropod Hydrobia ulvae

Laboratory microcosms were used to investigate the mud snail Hydrobia ulvae (Pennant) bioturbation activities and behavioural changes in response to snail density, algal food, sediment moisture content, light regime and water cover conditions. Density-dependent kinetics of bioturbated muddy areas were described by von Bertalanffy equations, which provided reliable estimates of mud surface covering rates by snail tracks (m² h⁻¹ snail⁻¹). Snails need a wet habitat to be active either covered by seawater or by moving in fluid layers for low-tide conditions. Light and microphytobenthic biomass, which are less potent to affect snail activity, are positively interrelated to increase covering rates in the tested chl a concentrations within the range of 1-15 μg g⁻¹. Experimental results suggested us the relevance of microphytobenthos migration processes in affecting crawling activities of H. ulvae that appeared to adjust their foraging efforts in response to benthic algal biomass. Behavioural processes of H. ulvae, in terms of floating, crawling, burrowing and inactive snails, were described using a Markov model. Finally, an empirical model based on von Bertalanffy equations was proposed to describe kinetics of sediment covering by snail tracks under the influences of snail density, sediment moisture content, chl a concentrations and the four combinations of presence/absence of light and seawater. This model should provide a base for further development of a hydrosedimentary model to simulate the effects of H. ulvae bioturbation activities on the resuspension of the intertidal cohesive sediment-water interface for various in situ conditions.     

Decomposition and nutrient release in halophytes of a Mediterranean salt marsh

This study dealt with the decomposition and nutrient release from the halophytes Atriplex portulacoides, Arthrocnemum macrostachyum, Limoniastrum monopetalum, and Spartina densiflora, the dominant species in the Castro Marim salt marsh, Portugal. Environmental effects on decomposition were also assessed. The study was carried out for one year using the in situ litterbag technique. S. densiflora showed a lower decomposition rate (k = 0.003 day⁻¹) than the other study species (k = 0.005-0.009). Study species showed similar decomposition patterns, that is, the weight loss mostly occurred during the autumn-winter period (study beginning in November). This indicates that temperature in this period did not hamper the decomposition process. The decomposition rate was positively affected by the initial N concentration (r² = 0.87, P < 0.05) and negatively by the C:N ratio (r² = 0.86, P < 0.05) in decomposing materials. At the end of the study, S. densiflora and L. monopetalum, the species with lower initial N concentrations, retained much higher proportion of initial N (89-109%) than the others (5-14%). Also, S. densiflora with the lowest P concentration retained higher proportion of initial P (48%) than the others (5-20%). Release of K and Mg were also slower from S. densiflora and was associated with their initial low concentration in this species. The lowest Mn release was observed from A. macrostachyum and also in relation to the lowest initial concentration. Our study supports the hypothesis that decomposition patterns of marsh species are mostly associated with differences regarding their morphology and chemical composition. Given the higher resistance of S. densiflora to decomposition, its progressive spreading may result in accumulation of organic detritus overtime in invaded salt marshes.     

Buffering role of the intertidal anemone Anthopleura aureoradiata in cercarial transmission from snails to crabs

In nature, parasite transmission from one host to the next takes place within complex biotic communities where non-host organisms can reduce transmission rates, for instance by preying on infective stages. We experimentally investigated the impact of four very different non-host organisms on the transmission of the microphallid trematode Maritrema novaezealandensis from its snail first intermediate host to its crustacean second intermediate host. We show that in laboratory mesocosms, accumulation of parasites in juvenile stalk-eyed mud crabs, Macrophthalmus hirtipes (Ocypodidae), was not reduced in the presence of cockles, Austrovenus stutchburyi, barnacles, Balanus sp., or the algae Enteromorpha spp., three organisms whose feeding mode or general abundance could negatively impact the parasite's infective stages (cercariae). In contrast, the presence of the anemone Anthopleura aureoradiata in the mesocosms caused a more than 4-fold reduction in the number of parasites acquired by crabs when compared to control mesocosms. Observations on fluorescent-dyed cercariae confirmed that they are ingested by anemones. Given the often high densities of anemones on mudflats, they may represent an important regulator of the abundance of M. novaezealandensis, and thus of the impact of this parasite on its hosts. These anemones may decrease cercarial transmission for many other trematode species as well. Our results stress the need for studies of parasite transmission in natural contexts rather than under simplified laboratory conditions.     

Macroalgal-mediated transfers of water column nitrogen to intertidal sediments and salt marsh plants

In many temperate estuaries, mats of opportunistic macroalgae accumulate on intertidal flats and in lower elevations of salt marshes, perhaps playing a role in linking water column nitrogen (N) supply to these benthic habitats. Using a flow-through seawater system and tidal simulator, we varied densities (equivalent to 0, 1, 2, or 3 kg m⁻² wet mass) of ¹⁵N-labelled macroalgae (Enteromorpha intestinalis) on estuarine sediments in microcosms with/without pickleweed (Salicornia virginica) to assess N transfers from algae. In the 6-week experiment, macroalgal biomass increased from initial levels in the lower density treatments but all algae lost N mass, probably through both leakage and decomposition. With all densities of algae added, sediments and pickleweed became enriched in ¹⁵N. With increasing mat density, losses of algal N mass increased, resulting in stepwise increases in ¹⁵N labeling of the deeper sediments and pickleweed. While we did not detect a growth response in pickleweed with macroalgal addition during the experiment, N losses from algal mats that persist over many months and/or recur each year could be important to the mineral nutrition of N-limited marsh plants. We conclude that N dynamics of intertidal sediments and lower salt marsh vegetation are linked to the N pools of co-occurring macroalgae and that further study is needed to assess the magnitude and importance of N transfers.     

Keeping warm in the cold: On the thermal benefits of aggregation behaviour in an intertidal ectotherm

Temperature is a primary determinant for species geographic ranges. In the context of global warming, most attention focuses upon the potential effects of heat stress on the future distribution of ectothermic species. Much less attention has, however, been given to cold thermal stress although it also sets species thermal window limits, hence distribution ranges. This study was conducted in winter on a South-Australian rocky shore in order to investigate the potential thermal benefits of the aggregation behavior observed in the dominant gastropod Nerita atramentosa. Thermal imaging was used to measure the body temperatures of 3681 aggregated individuals and 226 solitary individuals, and surrounding substratum temperature. N. atramentosa aggregates and solitary individuals were significantly warmer than their surrounding substratum. The temperature deviation between aggregates and substratum was, however, ca. 2 °C warmer than the one observed between solitary individuals and substratum. This result is critical since a body temperature increase of only a few degrees might enhance individual performance, hence organismal fitness, and could potentially drive changes in interspecific relationships. Besides, the potential higher thermal inertia of aggregates might increase the snail adaptive ability to abrupt environmental changes. We further investigate the potential thermal heterogeneity within an aggregate in order to identify any thermally advantageous position. Patch centers are significantly warmer than their edges, hence snails experience greater thermal advantages in the aggregate center. Finally, we examined the potential effect of aggregate size on snail temperature and thermal spatial heterogeneity. We identified an aggregate size threshold (216 individuals) beyond which all snails had equal thermal benefits, regardless of their spatial positions within an aggregate. While the determinism of this aggregate size threshold requires further investigations, the present work uniquely identified the thermal benefits of aggregation behavior for intertidal ectotherms under cold weather conditions. The implications of the present finding are discussed in the general framework of the ability of ectothermic populations to face environmental changes.     

Relative importance of fertiliser addition to plants and exclusion of predators for aphid growth in the field

Herbivore dynamics and community structure are influenced both by plant quality and the actions of natural enemies. A factorial experiment manipulating both higher and lower trophic levels was designed to explore the determinants of colony growth of the aphid Aphis jacobaeae, a specialist herbivore on ragwort Senecio jacobaea. Potential plant quality was manipulated by regular addition of NPK-fertiliser and predator pressure was reduced by interception traps; the experiment was carried out at two sites. The size and persistence of aphid colonies were measured. Fertiliser addition affected plant growth in only one site, but never had a measurable effect on aphid colony growth. In both habitats the action of insect predators dominated, imposing strong and negative effects on aphid colony performance. Ants were left unmanipulated in both sites and their performance on the aphid colonies did not significantly differ between sites or between treatments. Our results suggest that, at least for aphid herbivores on S. jacobaea, the action of generalist insect predators appears to be the dominant factor affecting colony performance and can under certain conditions even improve plant productivity.     

Water-borne cues from a shell-crushing predator induce a more massive shell in experimental populations of an intertidal snail

Inducible defenses are important in the life strategies of many taxa. In some species of marine gastropods, water-borne chemical cues from potential predators induce defensive changes in shell form and differences in growth rate. We examined such phenotypic plasticity in the direct-developing snail, Littorina subrotundata (Carpenter, 1864). Among experimental field populations of L. subrotundata exposed to differing intensities of predation by the purple shore crab, Hemigrapsus nudus (Dana, 1851), snails collected from predation-intense environments often had more massive shells than closely related snails from adjacent environments where predation was negligible. Snails collected from both environments were raised in tanks containing cages of H. nudus that were feeding on conspecific snails and compared to a control group raised in the absence of this stimulus. Most snails developed significantly more massive shells in the presence of the crabs suggesting that adaptive phenotypic plasticity may account for some of the variation we observed in the field. In one case, snails from a predation-intense environment did not exhibit a statistically significant amount of plasticity, but instead grew a more massive shell irrespective of the laboratory stimulus. We interpret this as evidence for a genetic difference in the plasticity of shell form among experimental populations, caused by intense selection by H. nudus. There was no statistical difference in the growth rates of snails among treatments.     

Metabolic heating in Mediterranean loggerhead sea turtle clutches

Offspring sex ratio is an important demographic parameter and, given its determination by incubation temperature in sea turtles, might be a key factor for their conservation under climate warming. An appealing approach to estimate hatchling sex ratios is to measure sand temperatures at nest depth and deduce hatchling sex ratios from a beforehand-established relationship of hatchling sex ratio and sand temperature. Such estimates will only be accurate though if metabolic heat produced by the embryos is considered. Judging whether metabolic heating has a potential effect on hatchling sex ratios without actually measuring temperature within clutches would greatly facilitate monitoring protocols. We tested for a relationship between the amount of metabolic heating and the number of developed embryos as well as clutch size in the largest known loggerhead sea turtle (Caretta caretta) population of the Mediterranean on Zakynthos (Greece). Temperatures were measured within 20 nests as well as at a reference site in the sand at nest depth. Metabolic heating was detected, but only during the last third of the incubation period did nests heat up considerably (1.6 °C on average) above the temperature of the surrounding sand. During the middle third of incubation, when sex is determined, the amount of metabolic heating was negligible. The amount of metabolic heating during the last third of the incubation duration was significantly correlated to the number of offspring developed to at least about 75% of incubation duration. This factor explained nearly 50% of variation in metabolic heating. Metabolic heating was also significantly correlated to clutch size. Given that clutch size within the Mediterranean is largest in Zakynthos loggerheads, we conclude that metabolic heating can be ignored in the estimate of hatchling sex ratios in Mediterranean loggerhead populations. These results thus provide the basis for a feasible monitoring of hatchling sex ratios in the loggerhead sea turtle in the Mediterranean.