Habitat selection in birds feeding on ocean shores: landscape effects are important in the choice of foraging sites by oystercatchers

Food availability is a fundamental determinant of habitat selection in animals, including shorebirds foraging on benthic invertebrates. However, the combination of dynamic habitats, patchy distributions at multiple spatial scales, and highly variable densities over time can make prey less predictable on ocean‐exposed sandy shores. This can, hypothetically, cause a mismatch between prey and consumer distributions in these high‐energy environments. Here we test this prediction by examining the occurrence of actively foraging pied oystercatchers (Haematopus longirostris) in relation to physical habitat attributes and macrobenthic prey assemblages on a 34 km long, high‐energy beach in Eastern Australia. We incorporate two spatial dimensions: (i) adjacent feeding and non‐feeding patches separated by 200 m and (ii) landscape regions with and without foraging birds separated by 2–17 km. There was no support for prey‐based or habitat‐based habitat choice at the smaller dimension, with birds being essentially randomly distributed at the local scale. Conversely, at the broader landscape dimension, the distribution of oystercatchers was driven by the density of their prey, but not by attributes of the physical beach environment. This scale‐dependence suggests that, on open‐coast beaches, landscape effects modulate how mobile predators respond to variations in prey availability.     

Spatial pattern of coastal fish assemblages in different habitats in North‐western Mediterranean

Variability of fish assemblages across habitat structures can depend on spatial scales. A hierarchical sampling design was used to assess the spatial variability of temperate fish assemblages in different habitats and at multiple scales. Underwater visual censuses were carried out along the coasts of Elba Island (NW Mediterranean) on Posidonia oceanica beds, rocky algal reefs and sandy habitat at three spatial scales, namely tens of metres (individual replicates), hundreds of metres (sites) and tens of kilometres (locations). At the assemblage level, there was a clear relationship between fish and habitat type and the observed habitat‐related differences were largely dependent on species identity. Fish assemblages on P. oceanica beds and rocky reefs shared a high number of species, whereas overlap with sandy assemblages was negligible. Multivariate analyses revealed significant differences in fish assemblages among habitats, although there was also a significant habitat × site interaction. These differences relied mainly upon assemblage composition and species richness. Assemblages on rocky reefs and P. oceanica meadows usually harboured a higher number of species and individuals compared with sandy assemblages. Nevertheless, the patterns of habitat‐related differences in species richness and, especially, in the total number of fish, changed significantly from site to site. Eight species showed significant differences over habitats, but they were not consistent due to the interaction of habitat with site. Predictability of fish at both assemblage and population levels decreased with the scale of observation, and the spatial pattern of fish observed at the smallest scale was likely dependent on factors other than habitat type.     

Inter-annual variability of North Sea plaice spawning habitat

Potential spawning habitat is defined as the area where environmental conditions are suitable for spawning to occur. Spawning adult data from the first quarter (January–March) of the International Bottom Trawl Survey have been used to study the inter-annual variability of the potential spawning habitat of North Sea plaice from 1980 to 2007. Generalised additive models (GAM) were used to create a model that related five environmental variables (depth, bottom temperature and salinity, seabed stress and sediment type) to presence–absence and abundance of spawning adults. Then, the habitat model was applied each year from 1970 to 2007 to predict inter-annual variability of the potential spawning habitat. Predicted responses obtained by GAM for each year were mapped using kriging. A hierarchical classification associated with a correspondence analysis was performed to cluster spawning suitable areas and to determine how they evolved across years. The potential spawning habitat was consistent with historical spawning ground locations described in the literature from eggs surveys. It was also found that the potential spawning habitat varied across years. Suitable areas were located in the southern part of the North Sea and along the eastern coast of England and Scotland in the eighties; they expanded further north from the nineties. Annual survey distributions did not show such northward expansion and remained located in the southern North Sea. This suggests that this species' actual spatial distribution remains stable against changing environmental conditions, and that the potential spawning habitat is not fully occupied. Changes in environmental conditions appear to remain within plaice environmental ranges, meaning that other factors may control the spatial distribution of plaice spawning habitat.     

Flatfish-habitat associations in Alaska nursery grounds: Use of continuous video records for multi-scale spatial analysis

Flatfish distributions have traditionally been described in terms of depth, temperature, and sediment characteristics, but other environmental variables may be important depending upon spatial scale. Surveys for age-0 northern rock sole (Lepidopsetta polyxystra) were conducted in five near-shore nursery sites at Kodiak Island, Alaska, using a towed camera sled integrated with navigational data. The continuous record of fish density and habitat features made possible a spatially comprehensive analysis of fish-habitat associations at several spatial scales, ranging from tens of kilometres to less than 1 m. A combination of multivariate statistical interpretation and geographic information systems (GIS) revealed that the distribution of juvenile rock sole was associated with environmental variables and spatial scales that are not normally detectable with usual flatfish— and habitat—sampling methods (i.e., trawls and grabs). Generalized additive models (GAM) incorporating habitat variables determined from video provided large improvements over models using only the traditional variables such as depth and sediment type. At the broadest (regional) scale of analysis, combinations of sediment composition, surface bedform, temperature, and density of worm tubes provided the best model for rock sole density. Within-nursery variation in fish density was modelled best with depth, habitat structural complexity created by emergent fauna and macroalgae, and worm tube density. At the microhabitat scale (< 1 m), there was little evidence of direct contact between rock sole and structures such as shell or algae. Rather, they were loosely associated on a scale tens of metres. This study showed that spatially comprehensive surveys can be conducted with towed camera systems and without the need for sediment grab samples. This approach yields detailed habitat information for fishes and the opportunity for landscape analysis of spatial patterns that will be important in conserving critical habitats for flatfishes and other fish species.     

Spatial distribution of crustaceans associated with shallow soft‐bottom habitats in a coral reef lagoon

The ecology and diversity of the shallow soft‐bottom areas adjacent to coral reefs are still poorly known. To date, the few studies conducted in these habitats dealing with macroinvertebrate fauna have focused on their abundance spatial patterns at high taxonomic levels. Thus, some aspects important to evaluate the importance and vulnerability of these habitats, such as species diversity or the degree of habitat specialization, have often been overlooked. In this study we compared the crustacean assemblages present in four different habitats at Magoodhoo Island coral reef lagoon (Maldives): coral rubble, sandy areas and two different seagrass species (Thalassia hemprichii and Cymodocea sp.). Forty‐two different crustacean species belonging to 30 families and four orders were found. ‘Site’ was a significant factor in all of the statistical analyses, indicating that tropical soft‐bottom habitats can be highly heterogeneous, even at a spatial scale between tens and hundreds of meters. Although traditionally it has been considered that seagrass beds host greater species diversity and abundance of organisms than adjacent unvegetated habitats, no differences in the univariate measures of fauna (abundance of organisms, number of species and Shannon diversity) were observed among habitats. However, sandy areas, coral rubble and seagrass beds exhibited different species composition of crustacean communities. The percentage of taxa considered as potential habitat specialists was 27% and the number of species exclusively occurring in one habitat was especially high in seagrass beds. Thus, degradation of this vegetated habitat would result in a great loss of biodiversity in tropical shallow soft‐bottom habitats.     

Spatial changes in a macrozoobenthic community along environmental gradients in a shallow brackish lagoon facing Sendai Bay, Japan

The spatial distribution, abundance, and assemblage structure of macrozoobenthos were examined at 45 stations in a brackish lagoon (Idoura Lagoon, Japan) to examine the animal–environmental relations in estuarine soft-bottom habitats. We found a total of 23 taxa; the polychaetes Heteromastus sp., Hediste spp., and Prionospio japonica and the isopod Cyathura muromiensis numerically dominated the community. Cluster analysis and one-way analysis of similarity (ANOSIM) identified seven groups of stations that had significantly different macrozoobenthic communities; these were subsequently consolidated into five habitat groups according to their association with environmental characteristics. Canonical correspondence analysis (CCA) showed that salinity, silt-clay content, and the oxidation–reduction potential (ORP) of the sediment strongly affected the macrozoobenthos distribution pattern in the lagoon, whereas other factors (e.g., relative elevation of the habitat and sediment organic content) had much weaker effects. Similarity percentages (SIMPER) procedures indicated that the polychaete Notomastus sp. and the bivalve Macoma contabulata were specific to habitats with low salinity and reduced mud, whereas the bivalve Nuttallia olivacea was specific to sandy bottoms. Heteromastus sp. and Hediste spp. achieved their highest densities in rather oxidized sediments. The acid-volatile sulfide (AVS) content in the sediment was suggested as another possible factor affecting macrozoobenthic density. Our results clearly demonstrate that macrozoobenthic assemblages in estuarine soft-bottoms have high spatial heterogeneity on a small scale (e.g., hundreds of meters) related to physical and chemical environmental changes. Our data also suggested the importance of sediment redox condition (e.g., ORP and AVS content) and sediment grain size as structuring factors in estuarine soft-bottom communities as well as the salinity in the habitat.     

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.     

Highly localised distribution patterns of juvenile sea cucumber Australostichopus mollis

Information on the environmental characteristics of the juvenile habitat of many deposit-feeding sea cucumber species is limited, despite most fished species exhibiting rapid localised depletion. The current study combined large and small scale surveying techniques within a New Zealand harbour to identify areas with high densities of juvenile Australostichopus mollis, a commercially valuable aspidochirote holothurian. Data from detailed surveys were used to relate densities of juveniles and adults with measures of physical habitat characteristics including depth, sediment facies type, grain size range, as well as measures of chlorophyll-a, phaeopigment, carbon and nitrogen content of surface sediment. Results revealed a highly localised distribution of juvenile A. mollis focused on one site associated with an area of high adult density. Sites of high juvenile A. mollis density were characterised by sediment qualities favouring epibenthic detritivorous deposit feeding, including high nitrogen content, high phaeopigment:chlorophyll-a ratio and small grain size. The high-density juvenile site had facies that were further characterised by the presence of large shell fragments (>10 cm length) of the horse mussel (Atrina zelandica), which may provide a unique settlement microhabitat for early juveniles. Unlike some other sea cucumber species, juvenile A. mollis shows no distinct spatial separation from adult sea cucumbers, no association with dense macroalgae and no clear preference for shallower depths than adults. Overall, the results illustrate the highly localised pattern of recruitment of this species to a widely distributed adult population, which may help to explain the lack of previous observations of juveniles in this species. These results indicate the importance of identifying and protecting what appear to be very specific juvenile habitats in deposit-feeding sea cucumbers to ensure continuing recruitment to exploited populations.     

Use of shallow water habitats by fish assemblages in a Mediterranean coastal lagoon

The present study investigates the different uses and the functional roles of shallow habitats for fish fauna in the Venice Lagoon, by applying the functional guilds approach. Temporal (seasons) and spatial (location) changes within the lagoon show different habitat uses by fish assemblages, not influenced by local factors. Unvegetated mud habitats (salt marsh creeks and sub-tidal mud flats) and sparsely vegetated (seagrass) habitats show a common nursery role, especially for marine migrant fishes; but, contrary to other similar areas elsewhere shallow seagrass beds in the Venice Lagoon do not have a primary nursery role. This latter habitat has a more important role as a spawning ground for a resident, highly specialized component of the fish community. The habitat uses and their role to fish fauna illustrate the balance between predation risk and foraging profitability, as major factors structuring the fish assemblages. Spatial effects at a larger scale also highlight general characteristics of the fish assemblages in the Venice Lagoon. These indicate differences between the Northern sub-basin and the other two lagoon sub-basins, due to differences in the hydrodynamic regime, habitats distribution, and contributions from land and sea.     

Spatial variability of the surf zone fish and macroinvertebrate community within dissipative sandy beaches in Oregon,USA

Habitat heterogeneity can influence biological communities by providing a diversity of areas that can be occupied by different species. Sandy beach surf zones are often considered homogenous environments; however, sand bars moved by currents and waves can produce trench‐like shapes or troughs that provide heterogeneity. The influence of habitat heterogeneity produced by sand movement is unclear despite the fact that surf zones are an important habitat for larval and juvenile fish and macrocrustaceans. To determine if, and how, the fish and macroinvertebrate communities present in trough and non‐trough or flat areas of Oregon surf zones differ, we compared species assemblages in both areas at three beaches adjacent to estuary mouths over 2 years. Troughs had different communities compared with flat areas, with higher total catch (mean ± SD = 123.2 ± 122.1 versus 43.6 ± 44.5 individuals × 100 m^?2) and taxon richness (6.7 ± 2.7 versus 4.0 ± 2.3 taxa); these differences were potentially due to water movement, prey availability and sediment size. The fish and macroinvertebrate communities did not vary between years but there were significant differences among beaches, with the most distinct community present at the only beach adjacent to an estuary without a jetty at its mouth, which was possibly due to higher species movement between the surf zones and estuary. Fish and macrocrustacean surf zone communities varied spatially within and among beaches in relation to habitat heterogeneity provided by sand movement and, potentially, the influence of adjacent habitats.     

Consistency of temporal and habitat-related differences among assemblages of fish in coastal lagoons

The consistency of habitat-related differences in coastal lagoon fish assemblages was assessed across different spatial and temporal scales. Multimesh gillnets were used to sample assemblages of fish on a monthly basis for 1-year in three habitats (shallow seagrass, shallow bare and deep substrata) at two locations (>1 km apart), in each of two coastal lagoons (approximately 500 km apart), in southeastern Australia. A total of 48 species was sampled with 34 species occurring in both lagoons and in all three habitats; species caught in only one lagoon or habitat occurred in low numbers. Ten species dominated assemblages and accounted for more than 83% of all individuals sampled. In both lagoons, assemblages in the deep habitat consistently differed to those in the shallow strata (regardless of habitat). Several species were caught more frequently or in larger numbers in the deep habitat. Assemblages in the two shallow habitats did not differ consistently and were dominated by the same species and sizes of fish, possibly due to habitat heterogeneity and the scale and method of sampling. Within each lagoon, very few between location differences in assemblages within each habitat were observed. Consistent differences in assemblages were detected between lagoons for the shallow bare and deep habitats, indicating there were some intrinsic differences in ichthyofauna between lagoons. Assemblages in spring differed to those in summer, which differed to those in winter for the shallow bare habitat in both lagoons, and the deep habitat in only one lagoon. Fish-habitat relationships are complex and differences in the fish fauna between habitats were often temporally inconsistent. This study highlights the need for greater testing of habitat relationships in space and time to assess the generality of observations and to identify the processes responsible for structuring assemblages.     

The influence of habitat availability on juvenile fish abundance in a northeastern New Zealand estuary

Management of fish populations is often focused on the exploitation of adult fish. Maintaining the habitat requirements of all life stages may also be an important consideration. We investigated the value of structured habitat to juvenile fishes within a northeastern New Zealand harbour using artificial seagrass units (ASUs). Specifically, we deployed ASUs across treatments with high vs. low habitat manipulations. We hypothesised that if the abundance of recruiting juvenile fishes was greater on the high habitat availability treatments this would suggest that the availability of habitat was limiting juvenile fish abundance. Our analyses were focused on the four most abundant fishes that settled on our ASUs: snapper (Pagrus auratus); spotty (Notolabrus celidotus); trevally (Pseudocaranx dentex); and an assemblage of pipefish species. For snapper, spotty and pipefish, abundance was greater on the high habitat availability treatments. This result would be unlikely to occur if the availability of juvenile fishes was limiting, suggesting that juvenile fish abundance is more likely to be limited by habitat. In terms of spatial gradients, spotty abundance and size increased with distance into the harbour. The spatial gradient present for spotty indicates the importance of placing the habitat dependency of juvenile fish into a landscape context. Overall, these results demonstrate that maintaining structured habitats may be an important consideration for some valuable inshore fish populations in northeastern New Zealand.     

Importance of feeding on regenerable parts of prey for juvenile stone flounder Platichthys bicoloratus in estuarine habitats

Prey availability is one of the most important factors affecting the quality of nursery grounds. Estuaries play an important role as nursery grounds for juvenile stone flounder, but the mechanism behind the consistently high availability of prey has never been examined. This study investigates which prey is mainly selected by juvenile stone flounder (15–55 mm standard length) in the estuary of the Natori River, northern Japan. In a cage experiment, juveniles showed positive selection for the palps of the spionid polychaete Pseudopolydora kempi in March, and for the siphons of the bivalve Nuttallia olivacea in April, May and June in both sandy and muddy-sand habitats. This selective predation showed that sublethal predation on regenerable parts of invertebrates is important for stone flounder. Nuttallia olivacea, the dominant bivalve in the estuary, was more abundant and in better somatic condition in the sandy area in spite of the stronger siphon-cropping pressure by juvenile stone flounder. These results confirm that sublethal predation on highly abundant benthos plays an important role in forming estuarine habitats into areas of high prey availability for juvenile stone flounder, which leads to their high growth rate.     

Spatial heterogeneity of epibenthos on artificial reefs: fouling communities in the early stages of colonization on an East Australian shipwreck

Artificial reefs are spatially complex habitats and serve as good model systems to study patterns of community succession and the response of epibiota to environmental clines over small spatial scales. Here, we quantified spatial heterogeneity in community composition and diversity of fouling communities across a number of environmental gradients that included water depth, surface orientation of habitats, exposure to currents, and shelter. Assemblage structure was quantified by spatially replicated photo transects on a recently scuttled large navy ship off the East Australian coast, lying in 27 m of water. A rich assemblage of epifauna had colonized the wreck within a year, dominated by barnacles, sponges and bryozoans. Community structure varied significantly over small spatial scales of meters to tens of meters. Depth, surface orientation and exposure were the major environmental drivers. Assemblages were substantially less diverse and abundant on the deepest (23 m near the seafloor) part of the hull with residual antifouling paint, on sheltered surfaces inside the wreck, and on the sediment‐laden horizontal surfaces. Overall, the wrecks’ habitat complexity corresponds with small‐scale heterogeneity in the fouling communities. This study supports the notion that wrecks enhance local diversity and biomass within the habitat mosaic of their location, and habitat complexity may be an important mechanism for this, as demonstrated by the large spatial variability in the assemblages documented here.     

Correlations between benthic habitats and demersal fish assemblages — A case study on the Dogger Bank (North Sea)

The interdependence between groundfish assemblages and habitat properties was investigated on the Dogger Bank in the North Sea. Abiotic habitat parameters considered included topography, hydrographic conditions, sediment composition, and the biotic habitat variable the prevailing benthic invertebrates. Distinct epi- and infauna communities occurred at different locations on the Dogger Bank. Fish assemblages were clearly linked to both the biotic and abiotic habitat characteristics.Overall, fish and benthic communities revealed similar spatial distribution, represented in the respective clusters of characteristic and abundant species. Distribution patterns corresponded with the prevailing abiotic conditions such as depth and sediment composition, which appear to relate to autecological preferences of individual species.The apparently most generalist species, grey gurnard (Eutrigla gurnardus) and dab (Limanda limanda) occurred at all stations and dominated in terms of biomass in most cases. The absolute numbers of grey gurnards were related to the abundance of suitable prey, invertebrate and fish species, which stomach analyses revealed as part of the diet in an independent study during the same research cruise. Haddock (Melanogrammus aeglefinus) and whiting (Merlangius merlangus) were only abundant at deep stations along the flanks of the bank. The occurrence of lemon sole (Microstomus kitt), American plaice (Hippoglossoides platessoides) and cod (Gadus morhua) was also positively correlated with depth, whereas especially lesser weever (Echiichthys vipera), sandeel species and solenette (Buglossidium luteum) occurred predominantly at the shallower sites. At the same time, individual fish species such as solenette and lesser weever were associated with high densities of selected epi- or infauna species.     

Regular habitat switch as an important feeding strategy of an opportunistic seabird species at the interface between land and sea

During deteriorated prey availability, purely pelagic, specialised seabird species have to alter their feeding strategy by extending foraging radii and/or time spent at sea or reducing feeding intervals of chicks. In contrast, more generalised species such as the opportunistic black-headed gull (Larus ridibundus) breeding at the German North Sea coast, can be assumed to react on prey shortages by switching foraging habitats. The coastal zone of the German North Sea provides a rich habitat mosaic consisting of the offshore zone, tidal flats and terrestrial habitats. Thus, we expected distinct temporal and spatial patterns of habitat switch in accordance with prey availability and environmental constraints. We carried out ship-based and aerial surveys as well as dietary analyses and observations on flight activity. We found a significant switch from terrestrial to marine feeding sites both on a daily basis (related to tidal cycle) and over the whole breeding season. Most likely, the latter switch is the result of lower prey availability in the terrestrial habitats and an increasing quality (in terms of prey abundance and energy intake) of the marine area. While there was only moderate variability in habitat use among different years, we revealed significant differences in the diet of birds from different colonies. The high dietary plasticity and flexible feeding strategy, switching between terrestrial and marine prey is certainly of major importance for the success of an opportunistic avian top predator in a complex coastal zone. It is suggested that – compared to situations elsewhere – the number of breeding pairs of black-headed gulls in the German North Sea coast are still stable due to the switch of foraging habitats performed by individuals in this region.     

Drift algae reduce foraging efficiency of juvenile flatfish

Although flatfish species utilise a wide range of habitats as adults, several species are confined to a very limited habitat as juveniles. Recruitment levels are dependent on the quality and quantity of these nursery areas and changes therein. In the Baltic Sea, these shallow environments are often subject to influxes of drifting macroalgae, which add structure to otherwise bare sandy substrate. Structure, such as vegetation, alters predator–prey interactions of a wide range of fauna and in an array of marine, freshwater, and terrestrial systems. The aim of our study was to assess the inhibition potential of drifting macroalgae on the foraging efficiency of juvenile flatfish (young of the year Scophthalmus maximus L., young of the year- and group 1 + Platichthys flesus L.) through a series of microcosm experiments. Our results show that foraging success is restricted by drift algae as predation efficiency of all predator species and size classes was negatively affected by the presence of macroalgae. Overall, there was a reduction in predation success by 80 ± 12% due to structural effects and/or the induced changes in water chemistry associated with the algae. Flatfish depend on shallow sandy areas as feeding and nursery grounds during their juvenile stage and frequently occurring macroalgal assemblages drastically change the features of the otherwise bare substrate, setting the stage for small-scale, localised processes potentially affecting population dynamics.     

Interannual and regional variability in distribution and ecology of juvenile pollock and their prey in frontal structures of the Bering Sea

The distribution, size, length-specific weight, growth, and feeding of age-0 walleye pollock (Theragra chalcogramma) were examined along with their prey distribution patterns in two contrasting transects over a 4-year period (1994–1997) in relation to biophysical properties of frontal regions around the Pribilof Islands, Bering Sea. There were significant interannual differences in catch of age-0 pollock, but transect and habitat differences (inshore vs. front vs. offshore) were not significant for either catch or size of pollock. There were significant variations in length-specific weight and growth of pollock, but the trends were inconsistent. Copepods dominated the zooplankton biomass in all habitats and years; there were no consistent differences in the densities of the dominant zooplankton taxa among the habitats. There were, however, strong habitat and transect differences in juvenile pollock diet, particularly for the larger and presumably rarer prey taxa (euphausiids, chaetognaths, fish). We did not find any evidence that occupying a particular habitat was beneficial to young pollock, although other factors (e.g. bioenergetic advantage and predation refuge) that we did not examine here could have been more variable and critical to pollock survival. In a physically dynamic system such as the Pribilof Islands, age-0 pollock may need to continuously search for optimal conditions of high prey availability and low predation pressure.