Seasonal Variation in the Covering ­Behaviour of the Echinoid Paracentrotus lividus (Lamarck)

Abstract. Covering behaviour is displayed by several regular echinoid species but its functional significance has yet to be quantified. Some of the most popular theories have interpreted this behaviour as an adaptive response to avoid over-exposure to light and/or as an anti-predator strategy. Since both predation pressure and light intensity may be seasonal as well as diurnal in nature, the aim of the present study was to quantitatively examine, in situ , the potential seasonality of covering behaviour in the purple sea urchin, Paracentrotus lividus (Lamarck). Data were collected over a one-year period at Lough Hyne, Co. Cork in Ireland. A peak in covering activity was observed over the summer months. Predator density and light intensity (sunlight hours) were also shown to vary seasonally at Lough Hyne. Covering behaviour in P. lividus , as in other echinoids, may be a response to a number of simultaneously interacting factors. At Lough Hyne, however, light intensity and covering item availability appear to be particularly important factors.     

Individual Variation in the Covering Behaviour of the Shallow Water Sea Urchin Paracentrotus lividus

Abstract. A number of recent field‐based studies have investigated the possible functions of covering behaviour in a range of echinoid species. Some of these have found important relationships between covering behaviour and environmental factors, such as light (in particular, ultra‐violet light) and covering item availability. However, these studies have often shown considerable within‐species variation, thus making covering functionality both within and between species difficult to interpret. The present study experimentally investigates individual variability in covering behaviour characteristics of the sea urchin Paracentrotus lividus (Lamarck). Data were collected bimonthly over a 1‐year period at Lough Hyne, Ireland, by observing P. lividus in outdoor aquaria. Significant variations in covering behaviour characteristics were observed as a function of individual size and mobility but not as a function of sex. A significant relationship between water temperature, mobility and covering behaviour was also found and is hypothesized to have implications for grazing activity in P. lividus. In contrast to previous field studies of P. lividus at Lough Hyne, no summer ‘peak’ in the proportion of individuals covering was observed. However, there was a significant relationship between local solar radiation and ‘time to first cover’. Overall, these data highlight (i) the ecological significance of individual variation in P. lividus covering behaviour and (ii) the importance of investigating multiple factors as potential determinants of echinoid covering behaviour.     

Stock Evaluation of Three Littoral Echinoid Species on the Catalan Coast North-Western Mediterranean

Abstract. We estimated the abundances of the three common littoral echinoid species Paracentrotus lividus, Arbacia lixula and Sphaerechinus granularis in a littoral zone between 0 and 10m depth on the Catalan shores NE Spain. Densities of each species were estimated from a total number of 11 571× 1 m² sampling units deployed at 152 sites along the 13 zones into which the coast was divided. The surface area available in each of these zones was estimated at the scale of the sampling unit 1 m² taking into account the fractal dimension of the shore. The stocks individuals > 2cm test diameter of these species for the whole Catalan shore ≅ 300 km, straight-line distance were estimated to be about 279.1 × 10⁶ individuals for P. lividus , 28.9 × 10⁶ individuals for A. lixula and 10.9 × 10⁶ individuals for S. granularis. These sea-urchins are key species in the regulation of algal communities and some of them P. lividus. S. granularis are of commercial interest. This information will provide a tool for the future monitoring and management of this resource, which now faces diverse pressures of human origin.     

Low densities of sea urchins influence the structure of algal assemblages in the western Mediterranean

Numerous studies of interactions between urchins and algae in temperate areas have shown an important structuring effect of sea urchin populations. These studies focused almost wholly on the effect of high urchin densities on laminarian forests. In contrast, algal communities below 5–6 m depth in the northwestern Mediterranean are characterised by low sea urchin densities (<5 ind m⁻²) and the absence of laminarian forests. No previous research has addressed sea urchin/algal interactions in this type of community. To determine the effect of the most abundant echinoid species, Paracentrotus lividus, on well-established algal communities in this area, we performed a removal–reintroduction experiment in rocky patches located between 13 and 16 m depth in the northwestern Mediterranean, where sea urchin densities ranged between 0.9 and 3.4 ind m⁻². After 6 months, the cover of non-crustose algae was significantly higher in the plots from which sea urchins had been removed than in control plots (84 vs 67% cover). These removal plots reverted to their original state upon reintroduction of sea urchins. The non-crustose algae consisted of turfing and frondose forms, with the former representing some 70% of the non-crustose algal cover. Change in the cover of turfing algae was responsible for the significant increase in algal development in the sea urchin removal plots. The response of frondose algae to the treatment varied between algal species. It is concluded that grazing by P. lividus exerts a significant effect on habitat structure, even in communities with low sea urchin densities, such as those found in vast areas of the Mediterranean sublittoral.     

Aspects of the ecology of a littoral chiton,Sypharochiton pellisekpentis (Mollusca: Polyplacophora)

On several Auckland shores, a littoral chiton, Sypharochiton pelliserpentis (Quoy and Gaimard, 1835), was widely distributed and common. At Castor Bay it was the commonest chiton, and its density equalled or exceeded that of the commonest limpet (Cellana spp.) over most of the inter‐tidal range. Spot measurements of population density were made at other sites including exposed and sheltered shores. The smallest animals were restricted to the lower shore in pools or on areas of rock which were slow to drain. Exclusive of these small animals, the population structure was similar in pools and water‐filled crevices situated either high or low on the shore. This was also true for populations of animals from drained surfaces, but, at both shore levels, the population structure in the two micro‐habitats (pools and drained surfaces) was distinct.

Single measurements of salinity in the very small volumes of water in which chitons were living gave a range of 13.3–45.8‰. In laboratory experiments at normal temperatures this species could survive these fluctuations for the maximum time possible between tides. Desiccation experiments, in which weight loss was expressed as a percentage of total water content, showed weight loss to depend on size. No significant differences in weight loss were found between populations from the two shore levels and the two micro‐habitats considered, though there were slight survival differences. Desiccation is probably an important factor influencing population structure, but loss of up to 75% of water content could be tolerated before 50% mortality occurred. Osmotic and desiccation stress evoked a characteristic behaviour pattern.     

Seasonal Changes in the Biochemical Composition of the Edible Sea Urchin Paracentrotus lividus Echinodermata: Echinoidea in a Lagoonal Environment

Abstract. Monthly soluble and insoluble protein, carbohydrate, lipid and ash levels and energetic content were measured in tissues of Paracentrotus lividus adults living in a lagoonal environment over a 1-year period to increase our understanding of resource allocation to the different tissues. The biochemical composition of P. lividus individuals in this environment is similar to that observed in other environments. The gonad and gut contained a high level of proteins soluble or insoluble, a lesser amount of lipids and, finally, a low level of carbohydrates. The main component of the test was ash, followed by protein levels. The lipid and carbohydrate levels in the test were very low. Seasonal variability in the biochemical composition of the different body parts was observed. In the gonad, an increase in the level of lipids was recorded concomitant to an increase in gonad weight. The lipids and carbohydrates, after an initial increase, decreased with the maturation of the gonad. The biochemical variations observed in the gut seem to be linked to both the reproductive cycle and the sea urchin's trophic activity. Variability in the test was low and irregular for all of the observed biochemical components. As P. lividus is an important member of the benthic community in the Urbinu lagoon, this echinoid represents an important store of materials and energy within this lagoonal environment.     

Comparing quality of estuarine and nearshore intertidal habitats for Carcinus maenas

Estuarine and nearshore marine areas are vital habitats for several fish and benthic invertebrates. The shore crab Carcinus maenas (Crustacea: Brachyura: Portunidae) inhabits a variety of coastal, estuarine and lagoon habitats. At low tide, habitat structural complexity may be most important for crabs in the intertidal, providing refuge from predation and desiccation. The quality of different vegetated and nonvegetated estuarine and rocky shore habitats in SW Portugal and SW England was evaluated for intertidal C. maenas populations. We estimated population density, size–structure, and potential growth (RNA/DNA ratios) to investigate habitat quality. Vegetated estuarine habitats supported higher crab densities, than nonvegetated estuarine and rocky shore habitats. Investigation of population size–structure revealed that all habitats seem important recruitment and nursery areas although estuarine habitats in SW Portugal appeared to support higher densities of new recruits than equivalent habitats in SW England. Significant variation was found in RNA/DNA ratios among habitats. Ratios were highest in the rocky shore suggesting a high quality habitat where growth potential is high. We speculate that competition from other top-predators (Pachygrapsus spp.) rather than low habitat quality may limit the occurrence of C. maenas in intertidal rocky shore habitats in SW Portugal. In estuarine environments RNA/DNA ratios were significantly higher in the vegetated than in the nonvegetated estuarine habitats in SW Portugal but not in SW England, suggesting geographic differences in the extent to which highly structure habitats represent high quality. Our results challenge the current paradigm that structured habitats are necessarily those of higher quality for C. maenas.     

Impacts of Habitat Complexity on Physiology: Purple Shore Crabs Tolerate Osmotic Stress for Shelter

Purple shore crabs, Hemigrapsus nudus (Crustacea: Decapoda), can survive indefinitely in salinities of 8 (25% SW), but also tolerate short-term exposure to salinities as low as 2. In the laboratory their salinity preference range, determined from choice chamber experiments, is 22–32 and they can discriminate between pairs of salinities separated by a difference of 2. These crabs show a strong positive thigmotaxis and a weak negative phototaxis and tend to choose environments with available shelter. The presence of shelter significantly alters the behaviour of this species. When shelter is available the salinity preference range is 10–32. Even in salinities below this preference range, the presence of shelter prolongs the time spent in the lower test salinities. This change in behaviour has implications on the crab's physiology: the haemolymph osmolality falls to lower levels when crabs remain in low salinity under shelters. In the field, H. nudus is found in creeks with salinities close to freshwater and they may remain in this salinity for up to 11 h, if there are rocks under which to shelter. An increase in habitat complexity increases the number of crabs that are found within the creek. These crabs in the low salinity environment have a lower haemolymph osmolality than crabs on the nearby open shore. In H. nudus the behavioural selection of a shelter appears to outweigh the physiological costs associated with osmotic regulation of the body fluids. Therefore, the distribution of H. nudus in estuaries may depend more on the availability of suitable habitats rather than the salinity tolerance of this species.     

Spatial variability of intertidal rocky shore assemblages in the northwest coast of Portugal

The spatial variability of rocky shore assemblages of the northwest Portuguese coast was studied in a total of 12 transects, visited twice between March and August 2003. Each transect was positioned from the upper to the lower shore and four replicate samples (50 × 50 cm) were taken at each visually identified assemblage. Multivariate analysis was used to test the null hypothesis of no significant differences among assemblages located on different heights along the transect and among assemblages located at the same height on different transects. The distribution pattern of organisms along the height was not consistent across transects and significant variability could be found in assemblages located at the same height on the shore. Globally, the variability from the lower to the upper shore (vertical axis) was larger than from transect to transect (horizontal axis) but the distribution pattern along height was clearer in sheltered transects than in exposed ones. The heterogeneity in composition between the visually identified assemblages, within the same transect, was, however, the major source of variability in the study area. In the rocky intertidal northwest coast of Portugal, where tidal amplitude is broad and extremely wave-exposed sites are scarce, height above chart datum was the most important factor determining the distribution and species composition of the assemblages. However, variability along the horizontal axis was also significant and should be considered in studies of spatial patterns of distribution of organisms within this area.     

Critical thermal maxima of common rocky intertidal fish and shrimps — A preliminary assessment

Rocky shore ecosystems are considered sentinels of climate warming because they are in close contact with the atmosphere and their shallow waters present low thermal inertia. Concerns on the vulnerability of rocky shore species subject to climate warming make the investigation of their thermal tolerance an urgent topic. The aim of this study was to determine the upper thermal limits of species that are common in tidal pools of rocky shore ecosystems of the Northeast Atlantic. The method used was the Critical Thermal Maximum (CTMax), which allowed the ranking of species in terms of their upper thermal limits as follows: Coryphoblennius galerita (32.0 °C), Palaemon serratus (33.0 °C), Gobius paganellus (33.1 °C), Palaemon elegans (33.4 °C), Lipophrys pholis (33.9 °C) and Paralipophrys trigloides (35.0 °C). Intraspecific variability was always lower than 2%.     

Characteristics of Habitats Used by Enteroctopus dofleini in Prince William Sound and Cook Inlet,Alaska

Abstract. Habitat characteristics associated with the giant Pacific octopus, Enteroctopus dofleini (Wülker 1910), were studied in Prince William Sound and Port Graham, Alaska, from beach walk, SCUBA and submersible surveys at depths of 0 to 197 m. Octopus counts on beach walk transects were positively correlated with soft substrates (sand, gravel or broken rubble), the presence of boulders, and dense kelp cover immediately offshore of the transect; and negatively correlated with depth on SCUBA transects. No significant habitat correlations were found with counts on submersible transects. On beach walks, octopus counts were reduced on hard substrates to 38 % of the counts on soft substrates. Counts increased five‐fold in the presence of boulders over counts in their absence, and increased fifteen‐fold adjacent to dense (> 75 %) kelp cover over counts adjacent to sparse (< 25 %) kelp cover. On SCUBA transects, the average density at less than 5 m depth was over five times that below 5 m. No trends in octopus size or sex ratio were detected with depth. Den use was inversely correlated with depth although there was no indication that den availability declined with depth. Octopuses were found at densities from 0 to 2.5 per 1000 m². These densities were only 1 to 50 % of densities of the same species recorded in British Columbia in the late 1970s and early 1980s. Few data are available to test recruitment, mortality, and habitat selection hypotheses that would account for differences between habitats. However, the presence of the highest octopus densities in intertidal and very shallow subtidal areas indicates the likely importance of near‐shore, shallow‐water habitats, and highlights the vulnerability of octopus populations to changes in these habitats.     

Strategies of marine dinoflagellate survival and some rules of assembly

Dinoflagellate ecology is based on multiple adaptive strategies and species having diverse habitat preferences. Nine types of mixing-irradiance-nutrient habitats selecting for specific marine dinoflagellate life-form types are recognised, with five rules of assembly proposed to govern bloom-species selection and community organisation within these habitats. Assembly is moulded around an abiotic template of light energy, nutrient supply and physical mixing in permutative combinations. Species selected will have one of three basic (C-, S-, R-) strategies: colonist species (C-) which predominate in chemically disturbed habitats; nutrient stress tolerant species (S-), and species (R-) tolerant of shear/stress forces in physically disturbed water masses. This organisational plan of three major habitat variables and three major adaptive strategies is termed the 3-3 plan. The bloom behaviour and habitat specialisation of dinoflagellates and diatoms are compared. Dinoflagellates behave as annual species, bloom soloists, are ecophysiologically diverse, and habitat specialists whose blooms tend to be monospecific. Diatoms behave as perennial species, guild members, are habitat cosmopolites, have a relatively uniform bloom strategy based on species-rich pools and exhibit limited habitat specialisation. Dinoflagellate bloom-species selection follows a taxonomic hierarchical pathway which progresses from phylogenetic to generic to species selection, and in that sequence. Each hierarchical taxonomic level has its own adaptive requirements subject to rules of assembly. Dinoflagellates would appear to be well suited to exploit marine habitats and to be competitive with other phylogenetic groups, yet fail to do so.     

Habitat provision of barnacle tests for overcrowded periwinkles

In habitats where competition for space is a shaping force of animal distribution such as in the intertidal rocky ecosystem, new habitats are readily taken by colonising species. We examined the importance of empty Chthamalus spp. tests as a habitat for the intertidal common periwinkle Melaraphe neritoides on Portuguese rocky shores. The role played by the space between neighbouring barnacles as a habitat for other species has been largely studied with regard to how an ecosystem functions, whereas the equivalent role of empty barnacle tests remains largely unknown. The small periwinkle is one of the most abundant snails in European rocky shores and is an important prey for key mobile predators. Biological facilitation is common in the rocky intertidal zone, where biological structures often potentiate the abundance of other species. The role played by barnacles as biological facilitators through habitat provision is not fully understood. In this study, the abundance and morphometric features of empty barnacle tests and their occupants were examined across shore levels and shores with differing exposures, as these are important gradients explaining barnacle distribution. The availability of empty barnacle tests was also experimentally manipulated to examine the percentage, time and length of occupation. Empty barnacles were more abundant on the midshore of sheltered shores and barnacle tests were wider on the upper shore but taller on the midshore. The minimum barnacle test occupation rate by the periwinkle was 70%. Barnacle shell height was an important factor determining snail occupation, where taller barnacles harboured significantly more, but not necessarily larger, snails. Snails outside the barnacle tests were significantly larger than those found within, suggesting that this crustacean group has an increased importance as a habitat for juvenile snails, thus potentially influencing the population dynamics of M. neritoides. We found that occupation of experimentally created empty barnacles was extremely high (70%) on the day after, and remained at 100% after 3 months of monitoring. Our study is the first of its kind to focus on the features of barnacle tests that snails occupy and their occupying snail traits. According to our results, it is clear that barnacles have an important role in providing additional habitat for young gastropods. The small periwinkle is an important grazer and prey for intertidal and subtidal foraging predators; hence, the potential refuge role of barnacle tests for juvenile M. neritoides may be important in the dynamics of intertidal communities.     

Fish assemblages in Tanzanian mangrove creek systems influenced by solar salt farm constructions

Deforestation of mangrove forests is common occurrence worldwide. We examined fish assemblage composition in three mangrove creek systems in Tanzania (East Africa), including two creeks where the upper parts were partly clear-cut of mangrove forest due to the construction of solar salt farms, and one creek with undisturbed mangrove forest. Fish were caught monthly for one year using a seine net (each haul covering 170 m²) within three locations in each creek, i.e. at the upper, intermediate and lower reaches. Density, biomass and species number of fish were lower in the upper deforested sites compared to the mangrove-fringed sites at the intermediate and lower parts in the two creeks affected by deforestation, whereas there were no differences among the three sites in the undisturbed mangrove creek system. In addition, multivariate analyses showed that the structure of fish assemblages varied between forested and clear-cut sites within the two disturbed creeks, but not within the undisturbed creek. Across the season, we found no significant differences except for a tendency of a minor increase in fish densities during the rainy season. At least 75% of the fishes were juveniles and of commercial interest for coastal fisheries and/or aquaculture. Mugil cephalus, Gerres oyena and Chanos chanos were the most abundant species in the forested sites. The dominant species in the clear-cut areas were M. cephalus and Elops machnata, which were both found in relatively low abundances compared to the undisturbed areas. The conversion of mangrove forests into solar salt farms not only altered fish assemblage composition, but also water and sediment conditions. In comparison with undisturbed areas, the clear-cut sites showed higher salinity, water temperature as well as organic matter and chlorophyll a in the sediments. Our results suggest that mangrove habitat loss and changes in environmental conditions caused by salt farm developments will decrease fish densities, biomass and species numbers as well as alter the overall fish assemblage composition in the salt farm area but not downstream in the creek.     

Influence of rock‐pool characteristics on the distribution and abundance of inter‐tidal fishes

Rock pools can be found in inter‐tidal marine environments worldwide; however, there have been few studies exploring what drives their, fish species composition, especially in Australia. The rock‐pool environment is highly dynamic and offers a unique natural laboratory to study the habitat choices, physiological limitations and adaptations of inter‐tidal fish species. In this study rock pools of the Sydney region were sampled to determine how the physical (volume, depth, rock cover and vertical position) and biological (algal cover and predator presence) parameters of pools influence fish distribution and abundance. A total of 27 fish species representing 14 families was observed in tide pools at the four study locations. The five most abundant species were Bathygobius cocosensis, Centropogon australis, Enneapterygius atrogulare, Lepidoblennius haplodactylus and Microcanthus strigatus, which together represented 71% of the total number of fish recorded. Larger rock pools containing more algal and rock ledge cover hosted a larger and more diverse population of fish. Furthermore, certain species were only found in pools with specific characteristics, such as the presence of loose shells, a variety algae or rock cover, suggesting a high degree of habitat specificity. By contrast, some species were ubiquitous and thus can likely tolerate a wide variety of physical conditions.     

Variations in crab claw morphology and diet across contrasting inter‐tidal habitats

Crabs are important predators of inter‐tidal ecosystems, controlling the abundance and distribution of their prey populations. Often the same crab species occupies several habitats and, although their effects on prey have been quantified across habitats, crabs’ dietary and morphological responses to differing environmental influences have been overlooked. Here, we used the crabs Eriphia verrucosa and Pachygrapsus marmoratus as model species to examine differences in claw morphometry – size and wear – and diet between rocky shore and heterogeneous sand flat habitats. We predicted that, intra‐specifically, crabs from rocky shores would consume more hard‐shelled prey owing to their high availability and consequently, would display chelipeds with the following claw characteristics: a higher degree of claw damage, stronger musculature (higher propel height) and increased mechanical advantage (defined as the ratio of input lever length to output lever length) than crabs in the heterogeneous sand flat habitats. Sampling was performed in heterogeneous sand flat habitats and rocky shores of the Central Portuguese coast. For each crab species, carapace width, diet composition and several claw morphometric measures were recorded, revealing significant intra‐specific differences (using multivariate analysis) between shore types. We found that E. verrucosa and P. marmoratus consumed more hard prey on rocky shore than on sand flat habitats, which resulted in rocky shore crabs having more accentuated dentition wear and larger musculature than their sand flat habitat counterparts. We suggest that the strong response of crab claw morphometry to environmentally induced diet variations is an important mechanism in the successful adaptation of crab species to inhabit differing habitats. A major implication is that the impact of the same species on prey may vary largely with habitat type as a result of predation efficiency varying with claw condition.     

Life-history traits displayed by neritic fish in the Benguela Current ecosystem

Three fish species, anchovy, hake and lanternfish, occupy different habitats in the neritic zone of the Benguela Current ecosystem. Compatibility between the predicted environmental variability in each habitat and the collection of life history traits found in each species is examined with simulation models. Variability in early-stage survival is introduced into the models by means of both random and non-random ("red") spectra. Traits exhibited by lanternfish are shown to be inefficient in filtering out random variability and the species therefore cannot prevent collapse in a non-randomly variable environment. By comparison, the traits found in anchovy and hake filter out part of both random and non-random variability in survival and allow a fairly stable population size to be maintained. It is argued that, as life-history traits are adapted to the spectrum of variability associated with the habitat, only anomalies with respect to this spectrum are likely to have a large influence on stock size.     

Ontogenetic habitat use by mangrove/seagrass-associated coral reef fishes shows flexibility in time and space

Tropical shallow-water habitats such as mangroves and seagrass beds are widely acknowledged as important juvenile habitats for various coral reef fish species, most of which are commercially important to fisheries. Spatio-temporal variability in ontogenetic habitat use by fish among these tropical coastal ecosystems has rarely been investigated, yet there are sufficient reasons to believe that this plays an important role. In the present study, we test the spatio-temporal variability in patterns of ontogenetic habitat use by some mangrove/seagrass-associated coral reef fishes (Lethrinus harak, Lethrinus lentjan, Lutjanus fulviflamma and Siganus sutor). Abundances of these four species were investigated during two years in Tanzanian coastal waters, using underwater visual census in mangrove, seagrass, shallow and deep mudflat, and shallow and deep coral reef habitats. The study covered four distinct seasons of the year and was done at two spatially separated (>40 km) locations. Averaged across locations, seasons and years, juveniles (≤10 cm length) of the four study species had significantly higher relative densities in shallow-water (mangroves and seagrass beds) than in deep-water habitats (deep mudflats or coral reefs), whereas the opposite pattern was found for the adults (>15 cm). These findings suggest a strong and general pattern of ontogenetic habitat shifts from shallow- to deep-water habitats. However, specific habitat-use patterns of juveniles as well as adults differed significantly in time and space. Various species showed subtle to considerable flexibility in juvenile as well as adult habitat use across seasons, years, or at different locations. Furthermore, for some species the data suggest presence of ontogenetic habitat shifts at one location but lack thereof at the other location. In summary, ontogenetic habitat use needs to be considered at various spatial and temporal scales for the interpretation of habitat utilization by fish during different life stages. This is important for conservation and management of these habitats, as essential habitats or seasons may be ignored or over-emphasized with respect to their importance for fish during different parts of their life cycle.     

Trophic relationships on a fucoid shore in south-western Iceland as revealed by stable isotope analyses, laboratory experiments, field observations and gut analyses

Rocky shores in the North Atlantic are known for their zonation patterns of both algae and animals, which can be expected to greatly affect food availability to consumers at different height levels on the shore. We tested the hypothesis that consumers would feed on the most abundant suitable food source in their surroundings. In total 36 species/taxa of common primary producers and consumers were sampled for stable isotope analyses from a sheltered fucoid shore at Hvassahraun in south-western Iceland. A selection of these species was also collected seasonally and from different height levels. Feeding experiments, field observations and gut analyses were also conducted. Our results were in good overall agreement with pre-existing knowledge of trophic relationships in the rocky intertidal. Consumers often appeared to be assimilating carbon and nitrogen from the most common diet in their immediate surroundings. The predator Nucella lapillus was thus feeding on different prey at different height levels in accordance with different densities of prey species. When tested in the laboratory, individuals taken from low on the shore would ignore the gastropod Littorina obtusata, uncommon at that height level, even when starved, while individuals from mid-shore readily ate the gastropod. This indicated that some kind of learned behaviour was involved. There were, however, important exceptions, most noteworthy the relatively small contribution to herbivores, both slow moving (the gastropod L. obtusata) and fast moving (the isopod Idotea granulosa and the amphipod Gammarus obtusatus) of the dominant alga at this site, Ascophyllum nodosum. The recent colonizer Fucus serratus seemed to be favoured. Selective feeding was indicated both by isotope signatures as well as by results of feeding experiments. Seasonal migrations of both slow and fast moving species could partly explain patterns observed.     

Genetic and morphological divergence in the purple sea urchin Paracentrotus lividus (Echinodermata, Echinoidea) across the African Mediterranean coast

The present investigation focuses on population structure analysis of the purple sea urchin Paracentrotus lividus across the African Mediterranean coast, with the main aim of assessing the influence of the Siculo-Tunisian Strait on gene flow disruption in this highly dispersive echinoid species. For this purpose, patterns of morphological and genetic variation were assessed among its populations from the western and eastern Mediterranean coasts. A total of 302 specimens from seven Tunisian sites were collected and examined for morphometric variability at twelve morphometric traits. Concordant results, inferred from CDA (canonical discriminant analyses), pairwise NPMANOVA (non parametric multivariate analysis of variance) comparisons and MDS (multidimensional scaling) plot, unveiled significant inter-population differences in the measured traits among the studied populations. Furthermore, the combined use of the one way ANOSIM (analysis of similarities) and the Discriminant/Hotelling analysis allowed unravelling two morphologically differentiated groups assigned to both western and eastern Mediterranean basins. The SIMPER (similarity percentages) routine analysis showed that total dry weight, test diameter and spine length were major contributors to the morphometric separation between locations and between groups. Pattern of phenotypic divergence discerned in P. lividus across the Siculo-Tunisian Strait is interestingly in congruence with that inferred from the genetic investigation of the purple sea urchin populations from the same region based on the analysis of the mtDNA COI (cytochrome oxidase I) gene in 314 specimens from nineteen locations covering a wider geographic transect, streching westward to the Algerian coast and eastward to the Libyan littoral. The specific haplotypic composition characterizing each Mediterranean basin, as inferred from the minimum spanning network, confirmed the geographic partioning of genetic variation, as revealed by F-statistics and AMOVA (analysis of molecular variance) analyses, yielding significant genetic differentiation between eastern and western Mediterranean populations. The newly detected phylogeographic patterns, observed for the first time in P. lividus throughout the explored distribution range, suggest the involvement of different biotic and abiotic processes in shaping such variation, and provide evidence that a large and geographically exhaustive dataset is necessary to unveil phylogeographic structure within widespread marine species, previously cathegorized as panmictic in part of their distribution range.