The first use of Fulton's K for assessing and comparing the conditions of inter‐tidal fish populations

Fulton's K condition factor was applied, for the first time, to inter‐tidal specimens of the shanny (Lipophrys pholis) and long‐spined scorpion fish (Taurulus bubalis) from two English rocky shore and two Welsh rocky shore sites during summer 2010 and winter 2011. As both species contribute to the diet of commercial species such as cod (Gadus morhua) and near‐threatened species such as the European otter (Lutra lutra), their condition may affect that of these predators. Fulton's K found that inter‐tidal Welsh fish maintained a ‘good’ condition between seasons, whereas the inter‐tidal English fish were in a poorer condition during winter. Although condition also changed amongst the sites on each coast, further studies are needed into fish morphologies, environmental parameters, prey availabilities and abundances, and fish specimen sex and maturities.     

Biotic consequences of a shift in invertebrate ecosystem engineers: Invasion of New Zealand rocky shores by a zone‐forming ascidian

The pervasive effects of invasive ecosystem engineers, that is those species that modify their environment, are well documented, but rarely have the broader impacts of one foundation invertebrate species being replaced by another been examined. In New Zealand, green‐lipped mussels, Perna canaliculus, commonly dominate wave‐exposed rocky shores. The recent appearance of an invasive ecosystem engineer, the ascidian Pyura doppelgangera, at the very northern tip of New Zealand now threatens to exclude these bivalves from this habitat. Here, we report major shifts in assemblages associated with the invader and chronicle its continued spread. We examined epibiota associated with clumps of mussels and clumps of Pyura from two rocky shore habitats—pools and emergent substrata at two locations. We detected some differences in species richness in biota associated with the two foundation species, but faunal abundance only differed between the locations. These minor changes were dwarfed by the shift in species composition within clumps of each foundation species. Molluscs, particularly gastropods, and crustaceans dominated the assemblage within mussels. In contrast, tubicolous polychaetes dominated the fauna associated with the ascidian. Sessile epifauna, notably barnacles and calcareous tube‐dwelling polychaetes, were common on mussels, but never encountered on the ascidian. Multivariate analysis revealed marked dissimilarity (>80%) between the characteristic mussel and ascidian faunas with virtually no overlap. This biotic shift overshadowed any differences between habitats and locations. The broader implications of these faunal shifts for local and regional patterns of biodiversity, as well as ecosystem function, remain unclear, but deserve further attention.     

Horizon scanning for alien predatory crabs: insights from South Africa

Due to the presence of few dominant predators on South African rocky shores, this coastline could be vulnerable to invasion by predatory crabs. This study applied horizon scanning to create an ordered watch list of alien crab species that could establish along this coastline under present-day and future temperature scenarios. This was done by: (i) identifying the species with both an invasion history and a possible pathway to South Africa; (ii) comparing the temperature ranges of the species’ native and introduced distributions to those of each of the four South African ecoregions; and (iii) ranking the species based on their potential ecological impacts. Of the 56 alien predatory crab species known worldwide, 28 species have pathways to South Africa. Incompatible temperature ranges excluded only two species from each ecoregion. Negative ecological impacts in their invaded ranges placed Japanese shore crab Hemigrapsus sanguineus, brush-clawed shore crab H. takanoi and Chinese mitten crab Eriocheir sinensis at the top of the watch list. This study highlights that many alien crab species have the potential to reach South Africa, with most likely to survive. This watch list should be used to support targeted monitoring and so facilitate early detection of these species, should they reach South Africa.     

Influence of tidal regime,diurnal phase,habitat and season on feeding of an intertidal crab

The relationship between diet and feeding activity of intertidal crabs, and environmental cycles (tidal, daily and seasonal), habitat and level of the intertidal zone (high/low) was studied using Neohelice granulata (Brachyura, Varunidae) as a model. This is a semi‐terrestrial burrowing crab occupying different habitats in the Southwestern Atlantic coasts and estuaries from bare low intertidal mudflats to high intertidal salt marshes, and from fine, organic matter rich sediment to very coarse sediment with low content of organic matter. The study was carried out in two contrasting habitats of three sites with diverse sets of physical and biological conditions. Diet and feeding of adult N. granulata were indirectly studied through the proportion of food items and the presence/absence of food in crab stomachs, respectively. This species has a dual mode of feeding: predominantly herbivorous (live plants or plant litter in salt marshes) or deposit feeder (superficial sediment and detritus in mudflats), but the quantity and quality of ingested food varies among habitats and sites. A trend to omnivory (including algae and conspecifics) was detected in relation to low quality of resources. Feeding activity modulated by a complex interaction of factors varied according to spatial and/or temporal changes in some natural cycles. Males and non‐ovigerous females fed preferably after dark and during submersion periods, but also after emersion periods if mudflat sediment remained wet; salt marsh crab feeding is somewhat independent of light and tidal cycles. Ovigerous females almost never fed. Both diet and feeding activity of this crab seem to be flexible traits adapted to different combinations of physical and biological factors.     

Spatio‐temporal variation in biomass of the deep‐sea red crab Chaceon affinis in Gran Canaria Island (Canary Islands,Eastern‐Central Atlantic)

The spatial and temporal biomass distribution of Chaceon affinis and its vulnerability to fishing activity in Gran Canaria (Canary Islands) were investigated. The first goal was to assess the influence of the slope steepness and substrate on the size of crab patches, size of the crabs, and crab biomass. The second goal was to evaluate spatial and temporal variation in the biomass over a 15‐month period. The last goal was to assess the influence of fishing activity upon the reduction in the biomass over the same 15‐month period. Only two or three locations in the sampling area generated high‐biomass contour patches. When these patches were superimposed on the isobath lines, they were coincident with the main depth range described for the species in the area. The map of the biomass values clearly showed three structures with cores of the highest biomass in both muddy and rocky‐muddy areas. The biomass was higher on muddy than on rocky‐muddy bottoms. Biomass was twice as high when steepness was reduced to one third between isobaths of 500 and 900 m. The size of crab patches increases linearly with the decrease in slope steepness. The spatial structure of crabs remained fairly stable over time, showing that biomass changes with depth over time. Maps of the estimated biomass values over the 15‐month period showed the same two main patches over time with the cores of highest biomass separated by a distance of between 4.2 and 4.5 km. Although the bathymetric distribution by sexes showed temporal changes, with a displacement to deeper areas made by both sexes over the studied period, only a partial temporal segregation between males and females was observed. During the study period, crabs underwent a significant decline in biomass and this was consistent with the combined catches of both commercial and experimental fishing in the area. Due to its low mobility, C. affinis is highly vulnerable to local depletion by intensive fishing efforts.     

Habitat modification in tidepools by bioeroding sea urchins and implications for fine‐scale community structure

By creating novel habitats, habitat‐modifying species can alter patterns of diversity and abundance in marine communities. Many sea urchins are important habitat modifiers in tropical and temperate systems. By eroding rocky substrata, urchins can create a mosaic of urchin‐sized cavities or pits separated by exposed, often flat surfaces. These microhabitats appear to harbor distinct assemblages of species. We investigated how a temperate rocky intertidal community uses three small‐scale (<100 cm²) microhabitats created by or adjacent to populations of the purple sea urchin (Strongylocentrotus purpuratus): pits occupied by urchins, unoccupied pits, and adjacent flat spaces. In tidepools, flat spaces harbored the highest percent cover of algae and sessile fauna, followed by empty pits and then occupied pits. The Shannon diversity and richness of these sessile taxa were significantly higher in flat spaces and empty pits than in occupied pits. The composition of these primary space holders in the microhabitats also varied. Unlike primary space holders, mobile fauna exhibited higher diversity and richness in empty pits than in flat spaces and occupied pits, although results were not significant. The protective empty pit microhabitat harbored the highest densities of most trophic functional groups. Herbivores, however, were densest in flat spaces, concordant with high algal coverage. These results suggest the habitats created by S. purpuratus in addition to its biological activities alter community structure at spatial scales finer than those typically considered for sea urchins.     

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.     

Shell use by sympatric hermit crab species in a seasonally open coastal lagoon in Northeastern Brazil

There are gaps in what is known about the patterns of gastropod shell use by hermit crabs in Brazilian lagoon areas, especially in Northeastern Brazil. However, this is important because the understanding of selection patterns provides information on life history and eco‐evolutionary conceptions of paguroids. The present study investigated the use of gastropod shells occupied by Pagurus criniticornis and Clibanarius sclopetarius in a coastal lagoon seasonally connected to an estuary and to the sea, correlating eco‐evolutionary aspects and hydrodynamic characters. The study was carried out between February 2013 and January 2014. Hermit crabs and their shells were identified and measured (hermit crabs’ cephalothorax shield length and width, major propodus length and height, weight, shell total length and width, shell aperture length and width). Partial least squares regression was used to analyse the morphometric data. Additionally, a multinomial proportions test was performed to infer patterns (inter‐/intra‐specific) of shells’ occupation. Hermit crab species occupied a total of 13 types of shells, predominantly those of Neritina virginea (67.83%). The shell weight was the most important determinant of shell occupancy in the morphometric model (variable importance in projection >1). The proportions of N. virginea shells used were similar in both species of hermit crabs, except between the sexes of P. criniticornis. Presumably, the high utilization rate of N. virginea shells is related to its abundance and dispersal, and to the shells’ suitability for hermit crabs, which is reflected by the morphometric model, as well as by the hydrodynamics of the lagoon‐estuarine environment. The relative occupation of different types of shells for each species of hermit crabs studied appears to be associated with regulation by inter‐/intra‐specific competition, which fosters the co‐occurrence of those populations. These results endorse coastal lagoons as a refuge and recruitment area for aquatic fauna. In addition, it highlights a challenge in the management and conservation of paguroid species whose population dynamics depend on the resources coming from the Gastropoda community.     

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.     

Marine invasive aliens on South African shores: implications for community structure and tropillc functioning

Up to 12 marine mollusc and three crab species are thought to have been deliberately or accidentally introduced to South Africa. Of these, only the Mediterranean mussel Mytilus galloprovincialis and the European shore crab Carcinus meanas have become invasive. M. galloprovincialis, probably introduced in the late 1970s, is already the dominant mussel throughout the Cape west coast. As a result, indigenous intertidal mussels Aulacomya ater have been largely displaced, mussel standing stock has increased and the upshore limit of mussel beds has become considerably elevated. Implications include competitive exclusion of large limpets but enhanced recruitment of juvenile limpets, increased habitat availability for mussel infauna, reduction of habitat for algal infauna and enhanced food availability for predators, particularly terrestrial species. C. maenas was first recorded from Table Bay Docks in 1983 and currently ranges from Camps Bay to Saldanha Bay. Although it is a voracious predator, it appears unable to colonize wave-swept shores, so is unlikely to displace indigenous crabs, or to severely impact prey species on the open coast. Valuable conservation areas and mariculture sites in sheltered lagoonal areas are, however, threatened by it.     

Hitching a ride on a hermit crabs home: Movement of gastropod shells inhabited by hermit crabs

There is little information on the movement and connectivity patterns for many species. The movement by shells occupied by the hermit crab Pagurus bernhardus and the organisms encrusting these shells was investigated on the south coast of Wales (UK). Hermit crabs shells moved considerable distances along the shore over 1 month periods, moving a minimum mean distance (±SE) of 148 (±8) and 174 (±9) m from release sites in the January and August respectively. Hermit crab-inhabited shells were also found to travel across habitats (sandy areas) that are unsuitable for both hermit crabs and epibionts. Through the examination of 150 of the most abundant shells used by P. bernhardus (Nucella lapillus), twelve epibiont species were found (10 animal and 2 algal species) and choice experiments demonstrated that hermit crabs preferred epibiont covered shells (84%), compared with bare shells (16%). The distance that shells were moved and the preference of hermit crabs for epibiont encrusted shells, coupled with the ability for epibionts to cross unsuitable habitats, may provide a dispersal advantage for epibiont species.     

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.     

Dietary analysis of harbour seals (Phoca vitulina) from faecal samples and overlap with fisheries in Erimo,Japan

The number of harbour seals in Japan has been rebounding since protection began in the mid‐1980s. With the increase in seal numbers, increased conflict with fisheries has occurred through depredation and the belief that seals compete with fisheries for prey. However, competition can only be determined if the prey species and quantities seals consume over time are known. We studied the diet of harbour seals in Erimo, site of the largest population of harbour seals in Japan, from 2011 to 2012 and assessed the degree of prey overlap with local fisheries. We used both hard parts and DNA techniques to identify prey items in seal scats, and compared these results to local fisheries data. A total of 46 prey occurrences was detected by both methods, of which 17 matched between techniques at least to the family level. Hard parts methods identified five incidences of prey undetected by DNA methods in five scats (one incidence per scat). DNA methods identified 24 additional prey occurrences in 13 scats, for which no hard part evidence for that prey had been found. This more than doubled the total number of prey occurrences across the 15 scats compared. Overall, the most frequently occurring harbour seal prey were walleye pollock (Theragra chalcogramma), sculpins and snailfishes. In contrast, the top three groups targeted by fisheries were codfishes, salmon and invertebrates. Many species common in the harbour seal's diet such as snailfishes and blennies were not targeted by fisheries. Fishes such as greenlings, sculpins, rockfishes and Japanese anchovy (Engraulis japonicus) were common in the diet of harbour seals, but made up a very small proportion of fisheries catches in Erimo. The importance of other prey species varied between seasons. Sculpins, greenlings and forage fishes were the top three prey groups for harbour seals in spring by percent modified frequency of occurrence, whereas the most important target groups by percentage mass caught by fisheries were codfishes, cephalopods and other invertebrates. Sculpins and rockfishes remained as important prey items for harbour seals in summer and autumn, in addition to codfishes, snailfishes and blennies. In contrast, the main groups caught by fisheries in summer were other fishes, particularly Japanese amberjack (Seriola quinqueradiata) and various species of sharks and eels, and invertebrates. By autumn, Erimo fisheries had focussed on catching salmon. Salmon were taken by harbour seals as well during this period, but at a relatively lower frequency compared to other prey groups. The results of our study show that although harbour seals consumed several of the prey species targeted by fisheries, the relative importance of these species to seals and fisheries and the seasons in which they were targeted were different.     

Localized versus regional adaptation in limpet shell morphology across the Iberian Peninsula

The climate‐envelope approach to predicting climate‐induced species range shift is limited. There are many possible reasons for this, but one novel explanation is that species adapt to changes in temperature at the expense of adaptation to other stressors. Here we test this hypothesis using the limpet Patella depressa (Mollusca, Patellidae), over a large geographical area covering most of the Atlantic coast of the Iberian Peninsula, known to consist of a genetically inter‐connected population. We examine limpet shell morphology on four shores in each of three regions, from Northern Spain to Southern Portugal. Within each region, shell morphology (measured as maximum shell profile to length ratio) varied between shore types differing in their insolation, wave action, microhabitat availability and biological factors. However, this ratio, which is known to be an adaptive response to heat stress, was found to be consistently higher in more southern latitudes despite differences between shore types being found in all regions. This implies that localized adaptation to shore type (most likely through phenotypic plasticity) is compromised by factors that change over latitudinal or regional scales, or which could occur in response to climate change. Although such climate‐induced changes may initially be localized, compromised adaptation (through phenotypic or genetic plasticity) may result in altered community interactions and potentially large shifts in community structure.     

Genetic differentiation of the soft shore barnacle Fistulobalanus albicostatus (Cirripedia: Thoracica: Balanomorpha) in the West Pacific

This study examined the phylogeography of the barnacle Fistulobalanus albicostatus, which inhabits mangroves and estuarine shores in the West Pacific. Differentiation in the mitochondrial cytochrome c oxidase subunit I (COI) and 12S ribosomal RNA (12S) genes of 401 specimens of F. albicostatus was examined in samples from 16 locations in the West Pacific, ranging from Honshu to Southern China. Our results revealed that F. albicostatus comprises two major clades exhibiting a COI divergence ranging from 1.25% to 2.8%. Clade A demonstrated the widest distribution, ranging from Japan to China, and was divided into three subclades occurring in the South China Sea (A1), Okinawa (A2), and Honshu, Korea and Qingdao (A3). Clade B was determined to be endemic to Okinawa; i.e. two endemic lineages occur in this island. Thus, F. albicostatus resembles several inter‐tidal species in having clades that are endemic to Okinawan waters. Nevertheless, in contrast to the rocky inter‐tidal barnacles Tetraclita spp. and Chthamalus malayensis, F. albicostatus was not found to be separated into continental and oceanic populations, but instead is divided into northern and southern clades, probably because of the Yangtze River discharge, which limits gene flow between the northern and southern populations.     

Gastropod shell repair tracks predator abundance

Predator‐prey interactions can shape biological communities and drive evolutionary change. The assessment of predation rates in modern and ancient ecosystems presents many challenges. We examined the utility of one common predation metric, shell repair frequency, as a proxy for crushing predation on marine gastropod populations. Our six localities near Bamfield, British Columbia, represented two contrasting predation regimes: three high‐wave‐energy (exposed) environments have low predation rates, while three low‐wave‐energy (protected) environments exhibit high predation rates. We measured the occurrence of repair scars for local populations of the gastropod Chlorostoma funebrale, and recorded multiple independent variables (shell measurements, water velocity, predatory crab abundance) at each locality. Fisher's exact test established that the repair frequencies observed in the protected localities were significantly greater than those of the exposed environments. Factors important in predicting the likelihood of an individual having a repair scar were examined using binomial logistic regression. A model containing the main effects of crab abundance, shell height, shell thickness and water velocity provided the best fit in predicting the presence of repair scars, with crab abundance having the largest contribution. The strong relationship between the presence of repair scars and predator abundance indicates that repair frequency is a valid tool for assessing predation intensity in gastropod populations.     

Evolution in Littorina: ecology matters

Organisms of marine rocky shores are exposed to physical stress from abiotic factors, such as temperature, salinity and wave action. These factors vary over compressed temporal and spatial scales, producing an exceedingly heterogeneous habitat with steep gradients of selection, and it seems likely that this has a strong influence on the evolution of populations of rocky shore organisms. With the periwinkles (genus Littorina) as a model group, I review strategies for coping with small-scale heterogeneous environments and what implications these strategies have on the evolution of these species.Some species of Littorina have long-lived pelagic larvae and sites of various habitats are thus recruited from a common gene pool. This largely prevents local adaptation but minor adjustments are possible through a plastic phenotype. Other species of the genus are directly developing with no larval dispersal and among these there is evidence of strong local adaptation forming distinct ecotypes in contrasting habitats by parallel evolution. In at least one of the directly developing species (L. saxatilis) divergent selection among ecotypes has resulted in partial reproductive barriers that further impede gene flow among ecotypes. Furthermore, convergent evolution among species has produced superficially similar morphs in different habitats. Ecotype formation, ecological reproductive barriers and convergence among species all indicate that ecological processes are critical for evolution of Littorina species.     

Effects of habitat complexity on cannibalism rates in European green crabs (Carcinus maenas Linnaeus, 1758)

The habitat in which predator–prey interactions take place may have a profound influence on the outcome of those interactions. Cannibalism is an intriguing form of predation whereby foraging by predators may contribute to the regulation of their own populations.This is particularly interesting in the case of invasive species, like the widely distributed European green crab (Carcinus maenas). This study explores how habitat complexity influences cannibalism rates in green crab populations of Prince Edward Island, Atlantic Canada. Both laboratory and field experiments were conducted to measure feeding rates by individual adult green crabs on a standard number of smaller conspecifics. In the laboratory, experimental treatments mimicked unstructured to increasingly structured habitats: water, sandy bottom, oyster shells, mussel shells, oyster shells with sandy bottom and mussel shells with sandy bottom. In those trials, adult green crabs consumed several times more juveniles on unstructured habitats than on the most structured ones, with a gradual decrease in predation rates across increasingly complex habitats. Field inclusion experiments used the same approach and were conducted in sandy bottoms, sandy bottoms with a layer of oyster shells and sandy bottoms with a layer of mussel shells. These trials showed similar patterns of decreasing feeding rates across increasingly complex habitats, but differences among treatments were not significant. These results support the idea that complex habitats have the potential to mediate predator–prey interactions, including adult–juvenile cannibalism in green crabs.     

Effect of crab size and habitat type on the locomotory activity of juvenile shore crabs, Carcinus maenas

Post-settlement processes are a major focus in the study of the dynamics of marine populations and communities. Post-settlement movement of juveniles is an important, but often ignored, process which affects local predator–prey and competitive interactions. We used benthic suction sampling and pitfall traps to examine density and locomotory activity of Carcinus maenas juveniles in different intertidal habitat types in the Rio Mira Estuary, Portugal, to better understand intra-specific interactions in a system where density-dependent processes are known to regulate population dynamics. As expected, significantly higher densities of juvenile shore crabs were found from bare mud compared to densely vegetated habitats. At the time of sampling, small and intermediate stages together outnumbered by far the larger juveniles. Conversely, larger crabs were much more frequent than smaller ones in traps. A locomotory index (LI), i.e. the ratio between crab catch in pitfall traps and their density within their moving range, is proposed as a measure of movement. LI analyses indicated that: (1) movement is an order of magnitude higher in large than small juveniles and much higher in sparse than dense vegetation cover; (2) activity of small juveniles is mostly crepuscular, regardless of vegetation cover; and (3) movement of large juveniles is very limited in dense Zostera patches, but very high in sparsely vegetated areas, during the day and night. These results suggest that small juveniles are relatively protected under dense vegetation cover due to lower mobility of larger crabs, and provide evidence of temporal segregation of activity windows between juvenile crabs of different sizes, which may be a key mechanism to reduce cannibalism and therefore increase the carrying capacity of nursery habitats.