Comparative studies reveal variability in the use of tidal stream environments by seabirds

The global increase in tidal stream turbine installations creates a need to identify and mitigate any impacts on seabird populations. Within Scotland, UK, the vulnerability of black guillemots Cepphus grylle and European shags Phalacrocorax aristotelis is dependent on their tendency to exploit microhabitats characterised by fast mean horizontal current speeds (≥2 ms^–1), and tidal states with maximum current speeds, within tidal stream environments. Identifying consistencies in their relative use of different microhabitats (fast versus slow mean horizontal current speeds) and tidal states (increasing/decreasing versus maximum currents) across these habitats could assist risk assessment and mitigation measures at both a regional and development site level. Datasets from shore-based surveys collated across 6 tidal stream environments showed that the probability of detecting foraging black guillemots and European shags tended to be higher in fast and slow microhabitats, respectively. However, differences between microhabitats were reversed and/or marginal in 3 out of the 5 sites used for each species. Differences between tidal states were almost always marginal. These variabilities show that a species' vulnerability could differ greatly among development sites, and environmental impact assessments (EIA) must quantify habitat-use using dedicated and site-specific surveys to reduce uncertainty. However, a greater understanding of the mechanisms underlying variation in the use of tidal stream environments is needed when selecting a suite of potential development sites that reduce the possibility of population-level impacts. The current collection of physical and biological data across tidal stream environments could therefore prove invaluable for the protection of seabird populations.     

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.     

Coupling bio-logging with nutritional geometry to reveal novel insights into the foraging behaviour of a plunge-diving marine predator

It is widely believed that predators maximise their energy intake while foraging and consume prey that are nutritionally similar. We combined GPS data loggers, miniaturised cameras, dietary sampling and nutritional geometry to examine the nutritional variability in the prey and selected diet, and foraging performance, of the masked booby (Sula dactylatra tasmani), a wild carnivore and marine top predator. Data loggers also revealed no significant differences between sexes in the foraging performance of chick-rearing adults. Females provided more food to their chicks than the males and, regardless of the nutritional variability of prey consumed, both sexes showed similar amounts of protein and lipid in their diets. Miniaturised cameras combined with nutritional analysis of prey provided, for the first time, fine-scale detail of the amounts of macronutrients consumed in each plunge dive and the overall foraging trip. Our methodology could be considered for future studies that aim to contribute to the general understanding of the behavioural and physiological mechanisms and ecological and evolutionary significance of animal foraging (e.g. energy expenditure budgets and prey selection for self- and offspring-feeding that could lead to sex-specific foraging strategies).     

Using at-sea experiments to study the effects of airguns on the foraging behavior of sperm whales in the Gulf of Mexico

Acoustic exposure and behavior of 8 sperm whales were recorded with acoustic and movement-recording tags before, during and after 5 separate 1–2 h controlled sound exposures of industry-provided airgun arrays. None of the 8 whales changed behavioral state (7 foraging, 1 resting) following the start of ramp-up at distances of 7–13 km, or full array exposures at 1–13 km. The most closely approached whale rested throughout exposure but started a foraging dive shortly after the airguns ceased, possibly indicating a delay in foraging during exposure. Using visual tracking and dead-reckoning of tagged animals, we found that their direction-of-movement was random with respect to the airguns, but correlated with their direction-of-movement just prior to the start of exposure, indicating that the tested whales did not show horizontal avoidance of the airguns. Oscillations in pitch generated by swimming movements during foraging dives were on average 6% lower during exposure than during the immediately following post-exposure period, with all 7 foraging whales exhibiting less pitching (p=0.014). Buzz rates, a proxy for attempts to capture prey, were 19% lower during exposure but given natural variation in buzz rates and the small numbers of whales, this effect was not statistically significant (P=0.141). Though additional studies are strongly recommended, these initial results indicate that sperm whales in the highly exposed Gulf of Mexico habitat do not exhibit avoidance reactions to airguns, but suggest they are affected at ranges well beyond those currently regulated due to more subtle effects on their foraging behavior.     

Seabird conservation and tidal stream and wave power generation: Information needs for predicting and managing potential impacts

Current development plans indicate that during the next decade there will be an increase in tidal stream and wave (TSW) power generation activity in Scottish Waters, together with the designation of additional offshore areas for seabird conservation. This paper summarises how TSW developments could affect seabirds, based on experience from other forms of disturbance, and explores the possible changes in behaviour and habitat that have the potential to increase a seabird's rate of energy acquisition (e.g. through enhancing prey abundance), or energy expenditure (e.g. through causing birds to commute further to find food, if they avoid foraging around developments placed in regular feeding areas). Changes to energy budgets could impact rates of reproduction and survival. Simulation modelling of seabird energetics and behaviour is one possible tool for predicting the direction and magnitude of population impacts caused by alterations to energy budgets, but is dependant on the availability and accuracy of necessary parameters. The later sections of the paper review the information needed for such models and shows that although some data are available regarding rates of energy expenditure during specific activities, more information is needed on seabird foraging rates. The paper also highlights how the susceptibility of a species to be impacted by future TSW development is likely to be related to their method of foraging, flight behaviour and ability to buffer against environmental fluctuations.     

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.     

Feeding preferences of estuarine mysids Neomysis integer and Rhopalophthalmus tartessicus in a temperate estuary (Guadalquivir Estuary,SW Spain)

Mysid shrimps are an important component of estuarine food webs because they play a key role in energy transfer as intermediate prey. We investigated the seasonal, tidal and depth specific variation in the diet of the estuarine mysids Neomysis integer and Rhopalophthalmus tartessicus and explored its implications for the planktonic community structure of a temperate estuary (Guadalquivir Estuary, SW Spain). Neomysis integer is an opportunistic omnivore feeding mainly on mesozooplankton and on members of the detrital–microbial loop, shifting prey seasonally according to availability. In contrast, R. tartessicus showed a more carnivorous diet and shifted its target prey during seasons of low resource availability. Despite statistically significant differences in diet composition, both species shared prey of similar size, particularly juvenile Mesopodopsis slabberi, the most abundant mysid species in this estuary, and copepods. Although these similarities imply inter-specific resource competition, their co-existence is achieved by niche partitioning and spatial segregation: the higher osmoregulatory capacity and foraging plasticity of N. integer confers a broader niche breadth for this species allowing N. integer to inhabit the more stressful oligohaline region of the estuary where R. tartessicus cannot survive. We propose that this mechanism relaxes the potential for competition between N. integer and R. tartessicus.     

Trophic ecology of European sardine Sardina pilchardus and European anchovy Engraulis encrasicolus in the Bay of Biscay (north-east Atlantic) inferred from δ13C and δ15N values of fish and identified mesozooplanktonic organisms

European sardine (Sardina pilchardus) and European anchovy (Engraulis encrasicolus) are two species of economical and ecological significance in the Bay of Biscay (north-east Atlantic). However, the trophic ecology of both species is still poorly known in the area, and more generally, few studies have considered the potential trophic overlap between sardines and anchovies worldwide. This study aims to highlight the trophic links between the mesozooplankton and adults of these two pelagic fish in the Bay of Biscay, through carbon and nitrogen stable isotope analysis (SIA). Mesozooplankton and individuals of sardines and anchovies were collected during one season (spring 2010), over spatially contrasted stations within the study area. First, the potential effect of preservation (ethanol vs. freezing) and of delipidation (by cyclohexane) on mesozooplankton δ¹³C and δ¹⁵N values was assessed. Results demonstrated the necessity to correct for the preservation effect and for lipid contents in mesozooplankton for further analyses of sardines' and anchovies' diet through SIA. Next, this study highlighted the interest of working on identified mesozooplanktonic organisms instead of undetermined assemblages when unravelling food sources of planktivorous fish using stable isotopes. The inter-specific variability of isotope values within a planktonic assemblage was effectively high, probably depending on the various feeding behaviours that can occur among mesozooplankton species. Intra-specific variability was also significant and related to the spatial variations of baseline signatures in the area. To investigate the foraging areas and potential diet overlap of S. pilchardus and E. encrasicolus, mixing models (SIAR) were applied. Both fish species appeared to feed mainly in the neritic waters of the Bay of Biscay in spring and to select mainly small- to medium-sized copepods (e.g. Acartia sp., Temora sp.). However, E. encrasicolus showed a greater trophic plasticity by foraging more offshore and on a wider range of prey sizes, while S. pilchardus seemed more limited to coastal areas and the mesozooplanktonic species of these waters for feeding.     

Influence of environmental and prey variables on low tide shorebird habitat use within the Robbins Passage wetlands,Northwest Tasmania

Shorebirds feed primarily on tidal flats, and their distribution over these flats is influenced by their prey and abiotic factors. These factors act by influencing the distribution and abundance of the prey, or the shorebirds ability to exploit it. The aims of this study were to investigate the low tide foraging distribution of shorebirds at four sites within the Robbins Passage wetlands, and the environmental and invertebrate factors that may influence their distribution. The greatest densities and number of shorebirds were found at Shipwreck Point and East Inlet. The shorebirds within-site distribution was also non-random, with the shorebirds present in greatest densities at the water's edge and low intertidal stratum, although this varied among species. Generally, on a small spatial scale, invertebrate diversity was positively correlated, and seagrass leaf mass was negatively correlated, with shorebird feeding density. On a large spatial scale, invertebrate biomass and seagrass root mass were positively correlated with shorebird feeding density. Invertebrate biomass and seagrass root mass explained 71% of the variance in total shorebird feeding density on the tidal flats. The variation in shorebird feeding density and diversity was therefore partly explained by invertebrate diversity and biomass, as well as the environmental factors seagrass roots and leaf mass and tidal flat area, although the strength of these relationships was influenced by the two different spatial scales of the study. The strength of the relationships between shorebird feeding density and the invertebrate and environmental variables was stronger on a large spatial scale. The presence of seagrass may have influenced shorebird-feeding density by affecting the invertebrate abundance and composition or the shorebirds ability to detect and capture their prey. The area of the tidal flat had opposing effects on the shorebird species. These results can be used to assist in the development of management plans for the Robbins Passage wetlands and the conservation of important shorebird areas.     

Modelling tidal stream power potential

In recent years, there has been an increase in interest in renewable energy sources for a number of reasons. A particular interest in tidal energy has developed within the UK due to its numerous sites of high current velocity. In this article a development, based upon previous work, of an existing hydrodynamic computational model is shown which is used to study the potential generation and the physical impacts of tidal stream farms. An idealised geometry is used to study the impacts of installed capacity and general layout of tidal stream farms and a realistic UK west coast model is used to examine the potential of presently proposed in-stream farms.     

The feeding ecology of the copepod Centropages typicus (Kröyer)

Here we report on the current knowledge on the feeding ecology of the planktonic copepod Centropages typicus. We describe the foraging process of C. typicus from the detection of prey to their digestion, considering also the effects of several physical variables on the feeding activity of the species. C. typicus is an omnivorous copepod that feeds on a wide spectrum of prey, from small algae (3–4 μm equivalent spherical diameter, ESD) to yolk-sac fish larvae (3.2–3.6 mm length). It uses both suspensivorous and ambush feeding strategies, depending on the characteristics of the prey. In general, C. typicus exhibits selection for large motile prey, such as ciliates or dinoflagellates, both in nature and laboratory, and this selective pattern is enhanced under moderate intensities of turbulence. Daily rations in the field are somewhat lower than those found in the laboratory, which indicates food limitation. This fact, together with the relatively modest capacity of the species to adapt to fluctuations in food availability may explain the geographical distribution of C. typicus, being restricted to near-shelf waters. In general, it does not seem that C. typicus feeding severely impacts planktonic populations. However, the occasional importance of the species in certain ecosystems is also apparent.     

Seasonal variation and tidal influences on estuarine use by bottlenose dolphins (Tursiops aduncus)

In order to show that dolphins use estuary habitats differently depending on the season and tidal state, possibly in response to prey distribution, temperature, risk of stranding and accessibility, Indo-Pacific bottlenose dolphins (Tursiops aduncus) were observed year-round during a 3-year study in the Clarence River estuary (CR) and Richmond River estuary (RR) in northern New South Wales, Australia. Peak dolphin sightings occurred during the spring season and one or 2 h prior to high tide. The spatial distribution of the dolphins in each estuary was analysed using the distance in kilometres that the dolphins travelled upstream with seasons and tidal phase as determinants. A General Linear Model showed that in the CR the dolphin spatial distribution in the estuary was not determined by season (F = 0.434, df = 3, P = 0.729) but was by tidal phase (F = 9.943, df = 3, P < 0.001) and the interaction between season and tidal phase (F = 3.398, df = 9, P < 0.002). However, in the RR the spatial distribution of the dolphin use of the estuary was not determined by either season (F = 1.647, df = 3, P = 0.194) or tidal phase (F = 0.302, df = 3, P = 0.824). In the CR, the spatial distribution of the dolphins was largest on high and flood tides. This pattern of spatial distribution may occur because the CR is a relatively shallow estuary and this increased spatial distribution may reflect a lower stranding risk and an increase in accessibility of shallow areas during periods of higher tide. These areas could also provide access to their preferred prey items of sea mullet (Mugil cephalus) and sand whiting (Sillago ciliata).     

潮流发电水轮机基于动量定理的性能计算方法研究

以潮流发电水轮机的设计为研究背景,分析和总结了基于动量定理方法的四种流管模型(单盘面-单流管模型、双盘面-单流管模型、单盘面-多流管模型和双盘面-多流管模型)在竖轴变攻角水轮机的水动力性能计算中的应用,特别是能量利用率预报方面的应用。说明基于动量定理的模型在求竖轴变攻角潮流发电水轮机的水动力性能方面,能够预报一些定性特性和总体趋势,为水轮机的设计和计算提供一种初步方法。     

Scale-dependent habitat use by a large free-ranging predator,the Mediterranean fin whale

Since the heterogeneity of oceanographic conditions drives abundance, distribution, and availability of prey, it is essential to understand how foraging predators interact with their dynamic environment at various spatial and temporal scales. We examined the spatio-temporal relationships between oceanographic features and abundance of fin whales (Balaenoptera physalus), the largest free-ranging predator in the Western Mediterranean Sea (WM), through two independent approaches. First, spatial modeling was used to estimate whale density, using waiting distance (the distance between detections) for fin whales along ferry routes across the WM, in relation to remotely sensed oceanographic parameters. At a large scale (basin and year), fin whales exhibited fidelity to the northern WM with a summer-aggregated and winter-dispersed pattern. At mesoscale (20–100 km), whales were found in colder, saltier (from an on-board system) and dynamic areas defined by steep altimetric and temperature gradients. Second, using an independent fin whale satellite tracking dataset, we showed that tracked whales were effectively preferentially located in favorable habitats, i.e. in areas of high predicted densities as identified by our previous model using oceanographic data contemporaneous to the tracking period. We suggest that the large-scale fidelity corresponds to temporally and spatially predictable habitat of whale favorite prey, the northern krill (Meganyctiphanes norvegica), while mesoscale relationships are likely to identify areas of high prey concentration and availability.     

Seasonal variations in prey selection by estuarine black-headed gulls (Larus ridibundus)

The number of black-headed gulls (Larus ridibundus) in the Clyde Estuary is large. In summer the average density has reached 1350 gulls km^?2 and in winter 180 gulls km^?2. This paper compares prey selection and feeding efficiency in gulls during summer and winter on tidal flats, and considers how seasonal differences may be adaptations to cope with seasonal changes in prey availability.Gross and net rates of energy intake were highest in summer because gulls captured more of the polychaete N. diversicolor than the amphipod C. volutator. In winter, gulls selected for C. volutator and therefore an energetically less profitable diet. Throughout the year gulls selected more C. volutator relative to N. diversicolor than expected on energetic grounds and so apparently did not maximize potential net rate of energy intake.Gulls used three techniques to capture prey and made most intensive use of the ‘crouch’ technique. Crouching gulls attained a much higher net rate of energy intake than ‘upright’ or ‘paddling’ gulls.A log-linear model showed that (a) season, water depth and gull density determined feeding technique and (b) feeding technique and season independently determined foraging success and prey selection. Thus gull density and water depth acted on prey selection through imposed variations in feeding technique.Reasons for gulls selecting energetically unprofitable C. volutator and for the use of several distinct feeding techniques are discussed.     

Small is profitable: No support for the optimal foraging theory in sea stars Asterias rubens foraging on the blue edible mussel Mytilus edulis

Doubt has been shed recently on the most popular optimal foraging theory stating that predators should maximize prey profitability, i.e., select that prey item that contains the highest energy content per handling time. We hypothesized that sea stars do not forage on blue mussels according to the classical optimal foraging theory but are actively avoiding damage that may be caused by e.g. capture of foraging on too-strong mussel shells, hence the sea stars will have a stronger preference for mussels that are smaller than the most profitable ones. Here we present experimental evidence of the sea star Asterias rubens as a predator that indeed chooses much smaller blue mussels Mytilus edulis to forage on than the most profitable ones. Hence this study does not support the optimal foraging theory. There may be other constraints involved in foraging than just optimizing energy intake, for example predators may also be concerned with preventing potential loss or damage of their foraging instruments.     

Elucidating cellular and behavioural effects of contaminant impact (polycyclic aromatic hydrocarbons,PAHs) in both laboratory-exposed and field-collected shore crabs,Carcinus maenas (Crustacea: Decapoda)

Monitoring effects of contaminants at lower levels of biological organisation (e.g. biochemical and cellular) allows for mechanistic evaluation of effects of contaminant exposure through laboratory exposures. However, higher level organism effects (e.g. physiological and behavioural) are deemed more ecologically-relevant. In the present study, cellular (cell viability and immune function), physiological (cardiac activity) and behavioural (foraging behaviour) responses were evaluated in field-collected shore crabs Carcinus maenas from three estuaries [a ‘relatively’ (polycyclic aromatic hydrocarbon) PAH-contaminated site (Plym Estuary) and two ‘comparatively clean’ field sites (Avon and Yealm Estuaries)] and compared with responses of crabs exposed in the laboratory to a model organic contaminant (PAH) pyrene (200 μg l⁻¹ for 28 days). The hypothesis that behavioural indicators may be more sensitive than other more traditional methods was tested. No significant impacts were observed at the cellular or physiological level in Plym-collected crabs (the ‘relatively’ contaminated site), but foraging behaviour was significantly altered (increased prey handling time) compared to individuals collected from the ‘relatively’ uncontaminated sites (Avon and Yealm). When given a cockle as a prey item, both Plym-collected and laboratory-exposed crabs took longer to handle and break into cockle shells. Therefore, ecologically-relevant behavioural observations may serve as valuableindicators of environmental quality.     

Prey versus substrate as determinants of habitat choice in a feeding shorebird

Many shorebirds on their non-breeding grounds feed on macrobenthic fauna which become available at low tide in coastal intertidal flats. The Eastern Curlew Numenius madagascariensis in Moreton Bay Australia, varies greatly in density among different tidal flats. This study asks: how important is the abundance of intertidal prey as a predictor of this variation? We quantified feeding curlews’ diet across 12 sites (different tidal flats, each re-visited at least eight times), through 970 focal observations. We also estimated the abundance of total macrobenthic fauna, potential prey taxa and crustacean prey on each tidal flat; measured as the number of individuals and a relative biomass index per unit substrate surface area obtained from substrate core samples. We estimated curlew density at each site using low-tide surveys from every site visit.     

水母伞径和饵料密度对海月水母(Aurelia sp.1)捕食率的影响

海月水母在全球范围内大量暴发,严重危害海洋生态系统的健康以及鱼类资源的可持续利用。本文以卤虫无节幼体为饵料,初步研究了水母伞径和饵料密度对海月水母个体捕食率的影响以及海月水母的昼夜捕食节律。研究结果表明,海月水母的伞径对其捕食率的影响极显著(P<0.01),在1—8cm范围内,随着伞径的增加个体捕食率线性增加,两者之间的关系式为y=0.044x+0.125(R2=0.9036)。饵料密度对海月水母捕食率的影响显著(P<0.05),在0.25—26.11mgC/L范围内,随饵料密度的增加个体捕食率呈线性增加趋势,关系式为y=0.508x(R2=0.958)。在饵料充足时,海月水母没有昼夜捕食节律。在高饵料密度下海月水母出现过剩捕食,但是捕食行为依然活跃。上述结果既为量化评估海月水母对浮游动物群体的捕食压力提供了基础参数,同时水母过剩捕食行为说明依据生物量和代谢率的推算结果可能低估自然海域水母队浮游动物的捕食压力。     

Prey selection and dietary overlap of shortfinned (Anguilla australis) and longfinned (A. dieffenbachii) eels during summer in the Horokiwi Stream,New Zealand

Benthic macroinvertebrates were collected and the diets of shortfinned eels (Anguilla australis) and longfinned eels (Anguilla dieffenbachii) inhabiting a small coastal New Zealand stream were examined to investigate intraspecific and interspecific trophic partitioning and prey selection. Our results showed intraspecific segregation of preferred prey among three size classes of juvenile shortfinned eels, but there was significant overlap between different size classes of longfinned eels. All size classes of both species of eel fed nocturnally on similar prey, and so there was no indication of temporal or trophic segregation. Therefore, habitat separation is assumed to be the main mechanism to reduce interspecific competition in these two co‐occurring species of eels.