Effect of disturbance on foraging: whelk activity on wave-exposed rocky shores with minimal tidal range

For slow-moving rocky intertidal predators such as whelks, frequent stress can be detrimental. Most studies on the foraging behavior of whelks have been conducted in regions where tidal amplitude is >1 m and on relatively small species. We studied the effects of wave action, desiccation, and water temperature on the foraging behavior of a large (approximately 80 mm) rocky shore whelk, Stramonita haemastoma, at the small-tide (<40 cm) but wave-exposed Israeli Mediterranean coast. We hypothesized that in such conditions foraging activity of this large predator will be greatly limited. Field observations show that when wave height exceeds 0.5 m, activity of whelks decreases by at least threefold relative to that seen under calm-sea conditions, and most whelks remain inside shelters. Wave height data from offshore moorings demonstrate that such conditions occur along the coast >85% of the time. Whelks move into shelters or completely disappear when exposed to prolonged desiccation conditions that can last for days along the shore during spring and autumn. Foraging activity is greater at night than in daytime and might be related to lighter winds and calmer sea conditions at night. Laboratory experiments show that elevated water temperatures during winter do not result in increased predation rates relative to those measured during spring. This indicates that a seasonal endogenous biological rhythm may exist. Our findings demonstrate that the foraging activity of large whelks can be extremely constrained by high rates of disturbance, that is, frequent wave action, and less frequent desiccation. We claim that high rates of disturbance in the eastern Mediterranean explain why large whelks are scarce in intertidal habitats where food is plentiful but suitable shelter is rare.Communicated by G.F. Humphrey, Sydney     

A morphological approach for relating decapod crustacean cephalothorax shape with distribution in the water column

Cephalothorax profiles for individuals of 18 decapod crustacean species were obtained from pictures of specimens captured using various experimental bottom trawls in the Mediterranean basin, at depths ranging from 200 to 4000 m in 2000 and 2002. Profiles were compared using the shape (outline) of the entire cephalothorax (including and excluding the rostrum). Principal component analysis (PCA) of the profiles, rostrum included, yielded two large species groupings related to rostrum size, species having a long rostrum being clustered on the positive portion of the first axis and species having a short rostrum being clustered on the negative portion of this same axis (the F1 axis explained 72.17% of the total variance). The PCA separated pelagic and endobenthic animals (with short rostrums) from nektobenthic animals (with long rostrums). Only the two deepest-dwelling species (dwelling at depths below 2000 m) were distanced from their respective groups. The pelagic shrimp Acanthephyra pelagica was clearly grouped with the nektobenthic species despite being reported to have a pelagic habitat. The nektobenthic deep-sea species Nematocarcinus exilis has a relatively medium-size rostrum and was placed among the pelagic–endobenthic species. The endobenthic and pelagic groups remained stable even when rostral morphometrics were excluded from the PCA analysis. The hypothesis that predation and diet are the fundamental determining factors for all habitats throughout most of the lifetimes of individuals and hence that species will evolve specialised phenotypes to adapt to local environmental conditions is considered in the discussion.Communicated by O. Kinne, Oldendorf/Luhe     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Nematocyst complements of nudibranchs in the genus <Emphasis Type="Italic">Flabellina</Emphasis> in the Gulf of Maine and the effect of diet manipulations on the cnidom of <Emphasis Type="Italic">Flabellina verrucosa</Emphasis>

Aeolid nudibranchs maintain functional nematocysts, which are sequestered from the nudibranchs’ cnidarian prey and provide protection against predators. Some species exhibit extensive variation in incorporated nematocysts, while others maintain a limited number of types. This study examines the apparent diversity in uptake and patterns of nematocyst incorporation among related species. Nematocyst complements were described for four Gulf of Maine nudibranch species in the genus Flabellina exhibiting a variety of feeding strategies and prey specificities. Diet manipulations were performed to examine the response to changing nematocyst availability using a generalist consumer, Flabellina verrucosa, to assess nematocyst uptake based on diet. The flabellinid species examined exhibited significant differences in nematocyst incorporation, reflecting differences in their specificity as predators and nematocyst types available in their natural prey. The nematocyst complement of F. verrucosa was the most variable and differed among collection regions. When diet was manipulated, nematocyst uptake depended on the prey the nudibranchs consumed, but when offered a variety of prey, F. verrucosa selectively preferred nematocysts from scyphistomae. The observed variation in nematocyst uptake among species and regions probably relates to environmental disparities among populations.     

Orientation of sandhoppers at different points along a dynamic shoreline in southern Tuscany

Orientation experiments were carried out on Talitrus saltator (Crustacea Amphipoda) at four points along 3 km on a dynamic sandy beach inside the Maremma Regional Park (Grosseto, Italy) to highlight behavioural variation related to distance from a river mouth, to erosion or accretion of shoreline, and to human trampling on the beach. Tests were performed using circular transparent Plexiglas arenas, contemporaneously at the four points. Replicates were made in 2 different months (September 2002 and May 2003), on 2–3 successive days, in the morning and afternoon. The distributions of the angles of orientation were compared for the different points and seasons, and multiple regression analysis was performed to test the effects of environmental and intrinsic variables on orientation. Sandhoppers showed the highest scatter at the eroded shoreline, intermediate scatter at the accreting beach most distant from the river mouth, and consistent orientation seaward at the least disturbed point. Orientation of sandhoppers was significantly affected by season, global radiation, time of day, distance from the river mouth, and human trampling. Sex and air humidity were of minor significance in the multiple regression model. The results, on the one hand, confirm plasticity in orientation of sandhoppers living on a dynamic shoreline, and on the other hand, show that variation in orientation could potentially be used as a bioindicator of shoreline changes.Communicated by R. Cattanevo-Vietti, Genova     

Concentrations of inorganic elements in bottled waters on the Swedish market

This study presents the concentrations of about 50 metals and ions in 33 different brands of bottled waters on the Swedish market. Ten of the brands showed calcium (Ca) concentrations ≤10 mg L⁻¹ and magnesium (Mg) levels <3 mg L⁻¹, implying very soft waters. Three of these waters had in addition low concentrations of sodium (Na; <7 mg L⁻¹), potassium (K; <3 mg L⁻¹) and bicarbonate (HCO₃ ≤31 mg L⁻¹). These brands were collected from barren districts. Nine of the brands were collected from limestone regions. They showed increased Ca-levels exceeding 50 mg L⁻¹ with a maximum of 289 mg L⁻¹. Corresponding Mg-levels were also raised in two brands exceeding 90 mg L⁻¹. Two soft and carbonated waters were supplemented with Na₂CO₃ and NaCl, resulting in high concentrations of Na (644 and 648 mg L⁻¹) and chloride (Cl; 204 and 219 mg L⁻¹). Such waters may make a substantial contribution to the daily intake of NaCl in high water consumers. The storage of carbonated drinking water in aluminum (Al) cans increased the Al-concentration to about 70 μg L⁻¹. Conclusion As there was a large variation in the material as regards concentrations of macro-elements such as Ca, Mg, Na, K and Cl. Supplementation with salts, e.g., Na₂CO₃, K₂ CO₃ and NaCl, can lead to increased concentrations of Na, K and Cl, as well as decreased ratios of Ca/Na and larger ratios of Na/K. Water with high concentrations of e.g., Ca and Mg, may make a substantial contribution to the daily intake of these elements in high water consumers. Al cans are less suited for storage of carbonated waters, as the lowered pH-values may dissolve Al. The levels of potentially toxic metals in the studied brands were generally low.     

Quantifying effects of pollution on biodiversity: a case study of highly diverse molluscan assemblages in the Mediterranean

Structured sampling designs are important in the assessment of environmental impacts of variable ecological systems. Recent developments have provided a useful framework extending existing univariate techniques into a multivariate context. Measures of taxonomic relatedness have also been introduced, which complement existing measures of diversity of assemblages. In this study, the potential effects of sewage discharge on spatial patterns of highly diverse molluscan assemblages in a Mediterranean rocky subtidal habitat were investigated. Nine 20 cm×20 cm quadrats were taken from each of three sites (80 m–100 m apart) within a putatively impacted location near a sewage outfall (I) and at each of two control locations (Cs) by destructive sampling by SCUBA divers at a depth of 3 m–4 m. A total of 5507 specimens of 151 species were collected. The average and the variance in total abundance of molluscs were greater, on average, at I than at Cs. Higher abundances at the sewage outfall were largely driven by greater numbers of juvenile molluscs. The Shannon diversity of molluscs (H′) was significantly lower at I, but no difference among locations was detected for the total number of species (S). In addition, the taxonomic distinctness (Δ*) of molluscs was greater at Cs, although it was more variable at I. Multivariate analyses showed that there was a significant difference in the structure of assemblages at I compared with Cs. The location near the outfall was characterized by greater abundances of several species, including especially the gastropods Pusillina philippi, Bittium latreilli, and Bittium reticulatum. There was also greater variability in the structure of assemblages among sites and among quadrats at control locations compared to those near the outfall. Using a suite of univariate and multivariate measures, including diversity indices, detailed information on taxonomic structure and analyses of variability at different spatial scales, provided useful insights into the effects of sewage impacts on these diverse assemblages. These results also highlighted the importance of analysing measures of variance, as well as mean in considering effects of stress in natural communities.     

The apportionment of quadratic entropy: a useful alternative for partitioning diversity in ecological data

Many methods that study the diversity within hierarchically structured populations have been developed in genetics. Among them, the analysis of molecular variance (AMOVA) (Excoffier et al., 1992) has the advantage of including evolutionary distances between individuals. AMOVA is a special case of a far more general statistical scheme produced by Rao (1982a; 1986) and called the apportionment of quadratic entropy (APQE). It links diversity and dissimilarity and allows the decomposition of diversity according to a given hierarchy. We apply this framework to ecological data showing that APQE may be very useful for studying diversity at various spatial scales. Moreover, the quadratic entropy has a critical advantage over usual diversity indices because it takes into account differences between species. Finally, the differences that can be incorporated in APQE may be either taxonomic or functional (biological traits), which may be of critical interest for ecologists.     

Quantitative analysis of aggregation behavior in juvenile blacktip sharks

Aggregation patterns of a population of juvenile blacktip sharks (Carcharhinus limbatus) within a summer nursery area were examined over a period of 3 years using an array of acoustic monitors. Individually coded acoustic tags were surgically implanted into 33–40 neonate sharks each year from 2000 to 2002 and their presence monitored continuously by the acoustic array. Data from the acoustic monitors was processed to estimate the center of activity location of each tagged shark every 30 min. Nearest neighbor analysis of shark locations revealed that sharks aggregated within the nursery in all years of the study. Sharks were aggregated most commonly during the early and late summer months (June, October and November) and became less common through the middle of the study period each year (July–September). Temporal periodicity within the data revealed a strong diel pattern with sharks aggregating during the day and dispersing at night, except in June when aggregations often persisted through the night. Aggregations were generally restricted to the mid and northern sections of the study site. Reasons for aggregations may include predator avoidance or improved feeding efficiency.