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Fisheries-induced mortality poses a threat to the conservation of small cetaceans, particularly those found in productive coastal waters. Harbour porpoises Phocoena phocoena are ubiquitous in UK seas, but despite their prevalence, knowledge gaps remain on fine-scale space use. A deeper understanding of how distributions vary in space and time is required for appropriate management. Citizen science programs which collect standardized data aboard platforms of opportunity can help uncover patterns and identify higher density areas in the time between surveys of absolute abundance. Here, we analyze citizen science data collected from ferries to investigate spatio-temporal patterns in porpoise densities along routes in UK waters. Region-specific detection functions and generalized additive models were used to estimate abundance and elucidate relationships between distributions and environmental variables. The highest densities during the study period (2006-2017) were found southwest of Cornwall, followed by the North Sea and the English Channel. Average density in the North Sea increased substantially during this time, suggesting new high-use areas along these routes. Density surface models showed strong relationships with coastal waters, sea surface temperature, chlorophyll, and water column dynamics. Contrasting preferences for stratified waters in the North Sea and well-mixed areas in the channel suggest distinct foraging behaviours. Higher density areas were identified in the southwestern and northeastern UK, indicating priority areas for conservation efforts, especially in Cornish waters where porpoises are highly vulnerable to bycatch. Findings highlight how citizen science, together with robust density estimation, can contribute to the conservation management of a common, although threatened, species.

Robust abundance estimates of wild animal populations are needed to inform management policies and are often obtained through mark–recapture (MR) studies. Visual methods are commonly used, which limits data collection to daylight hours and good weather conditions. Passive acoustic monitoring offers an alternative, particularly if acoustic cues are naturally produced and individually distinctive. Here we investigate the potential of using individually distinctive signature whistles in a MR framework and evaluate different components of study design. We analyzed signature whistles of common bottlenose dolphins, Tursiops truncatus, using data collected from static acoustic monitoring devices deployed in Walvis Bay, Namibia. Signature whistle types (SWTs) were identified using a bout analysis approach (SIGnature IDentification [SIGID]—Janik et al. 2013). We investigated spatial variation in capture by comparing 21 synchronized recording days across four sites, and temporal variation from 125 recording days at one high-use site (Aphrodite Beach). Despite dolphin vocalizations (i.e., echolocation clicks) being detected at each site, SWTs were not detected at all sites and there was high variability in capture rates among sites where SWTs were detected (range 0–21 SWTs detected). At Aphrodite Beach, 53 SWTs were captured over 6 months and discovery curves showed an initial increase in newly detected SWTs, approaching asymptote during the fourth month. A Huggins closed capture model constructed from SWT capture histories at Aphrodite Beach estimated a population of 54–68 individuals from acoustic detection, which overlaps with the known population size (54–76 individuals—Elwen et al. 2019). This study demonstrates the potential power of using signature whistles as proxies for individual occurrence and in MR abundance estimation, but also highlights challenges in using this approach.

Globally, reef manta rays (Mobula alfredi) are in decline and are particularly vulnerable to exploitation and disturbance at aggregation sites. Here, passive acoustic telemetry and a suite of advanced oceanographic technologies were used for the first time to investigate the fine‐scale (5‐min) influence of oceanographic drivers on the visitation patterns of 19 tagged M. alfredi to a feeding aggregation site at Egmont Atoll in the Chagos Archipelago. Boosted regression trees indicate that tag detection probability increased with the intrusion of cold‐water bores propagating up the atoll slope through the narrow lagoon inlet during flood tide, potentially transporting zooplankton from the thermocline. Tag detection probability also increased with warmer near‐surface temperature close to low tide, with near‐surface currents flowing offshore, and with high levels of backscatter (a proxy of zooplankton biomass). These combinations of processes support the proposition that zooplankton carried from the thermocline into the lagoon during the flood may be pumped back out through the narrow inlet during an ebb tide. These conditions provide temporally limited feeding opportunities for M. alfredi, which are tied on the tides. Results also provide some evidence of the presence of Langmuir Circulation, which transports and concentrates zooplankton, and may partly explain why M. alfredi occasionally remained at the feeding location for longer than that two hours. Identification of these correlations provides unique insight into the dynamic synthesis of fine‐scale oceanographic processes which are likely to influence the foraging ecology of M. alfredi at Egmont Atoll, and elsewhere throughout their range., A dynamic synthesis of fine‐scale oceanographic processes influences the foraging ecology of zooplanktivorous Mobula alfredi at Egmont Atoll in the Chagos Archipelago; British Indian Ocean Territory. These processes indicate that internal waves and cold‐water bores potentially transport zooplankton from the thermocline providing temporally limited feeding opportunities which are enhanced by the tide. Furthermore, processes such as Langmuir Circulation may provide extended foraging opportunities by transporting and concentrating zooplankton in the region.

Given the recent trend towards establishing very large marine protected areas (MPAs) and the high potential of these to contribute to global conservation targets, we review outcomes of the last decade of marine conservation research in the British Indian Ocean Territory (BIOT), one of the largest MPAs in the world. The BIOT MPA consists of the atolls of the Chagos Archipelago, interspersed with and surrounded by deep oceanic waters. Islands around the atoll rims serve as nesting grounds for sea birds. Extensive and diverse shallow and mesophotic reef habitats provide essential habitat and feeding grounds for all marine life, and the absence of local human impacts may improve recovery after coral bleaching events. Census data have shown recent increases in the abundance of sea turtles, high numbers of nesting seabirds and high fish abundance, at least some of which is linked to the lack of recent harvesting. For example, across the archipelago the annual number of green turtle clutches (Chelonia mydas) is ~ 20,500 and increasing and the number of seabirds is ~ 1 million. Animal tracking studies have shown that some taxa breed and/or forage consistently within the MPA (e.g. some reef fishes, elasmobranchs and seabirds), suggesting the MPA has the potential to provide long-term protection. In contrast, post-nesting green turtles travel up to 4000 km to distant foraging sites, so the protected beaches in the Chagos Archipelago provide a nesting sanctuary for individuals that forage across an ocean basin and several geopolitical borders. Surveys using divers and underwater video systems show high habitat diversity and abundant marine life on all trophic levels. For example, coral cover can be as high as 40–50%. Ecological studies are shedding light on how remote ecosystems function, connect to each other and respond to climate-driven stressors compared to other locations that are more locally impacted. However, important threats to this MPA have been identified, particularly global heating events, and Illegal, Unreported and Unregulated (IUU) fishing activity, which considerably impact both reef and pelagic fishes.

Funding: D.J.F.R. and D.T. are supported by National Capability funding from NERC to SMRU (Grant No. SMRU1001). There is high spatial overlap between grey seals and shipping traffic, and the functional hearing range of grey seals indicates sensitivity to underwater noise emitted by ships. However, there is still very little data regarding the exposure of grey seals to shipping noise, constraining effective policy decisions. Particularly, there are few predictions that consider the at-sea movement of seals. Consequently, this study aimed to predict the exposure of adult grey seals and pups to shipping noise along a three-dimensional movement track, and assess the influence of shipping characteristics on sound exposure levels. Using ship location data, a ship source model, and the acoustic propagation model, RAMSurf, this study estimated weighted 24-h sound exposure levels (10–1000 Hz) (SELw). Median predicted 24-h SELw was 128 and 142 dB re 1 μPa2s for the pups and adults, respectively. The predicted exposure of seals to shipping noise did not exceed best evidence thresholds for temporary threshold shift. Exposure was mediated by the number of ships, ship source level, the distance between seals and ships, and the at-sea behaviour of the seals. The results can inform regulatory planning related to anthropogenic pressures on seal populations. Postprint

Funding: Natural Environment Research Council (NERC), and the Department for Business, Energy and Industrial Strategy (BEIS; project no. OESEA-16-76/OESEA-16-78) (MIDC). National Capability funding from NERC to SMRU (grant no. SMRU10001) (DFJR & DT). Rapid development of a successful foraging strategy is critical for juvenile survival, especially for naïve animals that receive no parental guidance. However, this process is poorly understood for many species. Although observation of early-life movements is increasingly possible with miniaturisation of animalborne telemetry devices, analytical limitations remain. Here, we tracked 29 recently-weaned, grey seal Halichoerus grypus pups from colonies in two geographically distinct regions of the United Kingdom. We analysed at-sea movements of pups throughout their initial months of nutritional independence to investigate the ontogeny of behaviour-specific (foraging and travelling) movement patterns. Using generalized hidden Markov models (HMMs), we extended the conventional HMM framework to account for temporal changes in putative foraging and travelling movement characteristics, and investigate the effects of intrinsic (sex) and extrinsic (environment) factors on this process. Putative foraging behaviour became more tortuous with time, and travelling became faster and more directed, suggesting a reduction in search scale and an increase in travel efficiency as pups shifted from exploration to an adult-like repeatable foraging strategy. Sex differences in movement characteristics were evident from colony departure, but sex-specific activity budgets were only detected in one region. We show that sex-specific behavioural strategies emerge before sexual size dimorphism in grey seals, and suggest that this phenomenon may occur in other long-lived species. Our results also indicate that environmental variation may affect the emergence of sex-specific foraging behaviour, highlighting the need to consider interacting intrinsic and extrinsic factors in shaping movement strategies of long-lived vertebrates. Moreover, comparing the behavioural state estimations to those of a conventional HMM (no variation in state-specific movement parameters) revealed differences in the amount and location of foraging activity, with implications for spatial conservation management. Overlooking intrinsic and extrinsic variation in movement processes could distort our understanding of foraging ecology, population dynamics, and conservation requirements. Postprint

Mid-latitude (∼30-60°) seasonally stratifying shelf-seas support a high abundance and diversity of marine predators such as marine mammals and seabirds. However, anthropogenic activities and climate change impacts are driving changes in the distributions and population dynamics of these animals, with negative consequences for ecosystem functioning. Across mid-latitude shelf-seas marine mammals and seabirds are known to forage across a number of oceanographic habitats that structure the spatio-temporal distributions of prey fields. Knowledge of these and the bio-physical mechanisms driving such associations are needed to improve marine management and policy. Here, we provide a concise and easily accessible guide for both researchers and managers of marine systems on the predominant oceanographic habitats that are favoured for foraging by marine mammals and seabirds across mid-latitude shelf-seas. We (1) identify and describe key discrete physical features present across the continental shelf, working inshore from the shelf-edge to the shore line, (2) provide an overview of findings relating to associations between these habitats and marine mammals and seabirds, (3) identify areas for future research and (4) discuss the relevance of such information to conservation management. We show that oceanographic features preferentially foraged at by marine mammals and seabirds include shelf-edge fronts, upwelling and tidal-mixing fronts, offshore banks and internal waves, regions of stratification, and topographically complex coastal areas subject to strong tidal flow. Whilst associations were variable across taxa and through space and time, in the majority of cases interactions between bathymetry and tidal currents appear to play a dominant role, alongside patterns in seasonal stratification and shelf-edge upwelling. We suggest that the ecological significance of these bio-physical structures stems from a capacity to alter the densities, distributions (both horizontally and vertically) and/or behaviours of prey in a persistent and/or predictable manner that increases accessibility for predators, and likely enhances foraging efficiency. Future conservation management should aim to preserve and protect these habitats. This will require adaptive and holistic strategies that are specifically tailored to the characteristics of an oceanographic feature, and where necessary, evolve through space and time in response to spatio-temporal variability. Improved monitoring of animal movements and bio-physical conditions across shelf-seas would aid in this. Areas for future research include multi-disciplinary/trophic studies of the mechanisms linking bio-physical processes, prey and marine mammals and seabirds (which may elucidate the importance of lesser studied features such as bottom fronts and Langmuir circulation cells), alongside a better understanding of how predators perceive their environment and develop foraging strategies during immature/juvenile stages. Estimates of the importance of oceanographic habitat features at a population level should also be obtained. Such information is vital to ensuring the future health of these complex ecosystems, and can be used to assess how anthropogenic activities and future environmental changes will impact the functioning and spatio-temporal dynamics of these bio-physical features and their use by marine predators.

Underwater noise pollution from shipping is a significant ecological concern. Acoustic propagation models are essential to predict noise levels and inform management activities to safeguard ecosystems. However, these models can be computationally expensive to execute. To increase computational efficiency, ships are spatially partitioned using grids but the cell size is often arbitrary. This work presents an adaptive grid where cell size varies with distance from the receiver to increase computational efficiency and accuracy. For a case study in the Celtic Sea, the adaptive grid represented a 2 to 5 fold increase in computational efficiency in August and December respectively, compared to a high resolution 1 km grid. A 5 km grid increased computational efficiency 5 fold again. However, over the first 25 km, the 5 km grid produced errors up to 13.8 dB compared to the 1 km grid, whereas, the adaptive grid generated errors of less than 0.5 dB.

Shelf-seas are highly dynamic and oceanographically complex environments, which likely influences the spatio-temporal distributions of marine megafauna such as marine mammals. As such, understanding natural patterns in habitat use by these animals is essential when attempting to ascertain and assess the impacts of anthropogenically induced disturbances, such as those associated with marine renewable energy installations (MREIs). This study uses a five year (2009–2013) passive acoustics (C-POD) dataset to examine the use of an oceanographically dynamic marine renewable energy test site by small cetaceans, dolphins (unspecified delphinids) and harbour porpoises Phocoena phocoena, in the southern Celtic Sea. To examine how temporal patterns in habitat use across the site related to oceanographic changes occurring over broad seasonal scales as well as those driven by fine scale (bi-weekly) localised processes (that may be masked by seasonal trends), separate analyses were conducted using (1) all daily animal detection rates spanning the entire five year dataset and (2) daily animal detection rates taken only during the summer months (defined as mid-June to mid-October) of 2010 (when continuous monitoring was carried out at multiple discrete locations across the site). In both instances, generalised additive mixed effects models (GAMMs) were used to link detection rates to a suite of environmental variables representative of the oceanography of the region. We show that increased harbour porpoise detection rates in the late winter/early spring (January–March) are associated with low sea surface temperatures (SST), whilst peaks in dolphin detection rates in the summer (July–September) coincide with increased SSTs and the presence of a tidal-mixing front. Moreover, across the summer months of 2010, dolphin detection rates were found to respond to small scale changes in SST and position in the spring-neap cycle, possibly reflective of a preference for the stratified waters immediately offshore of the front. Together, these findings suggest that habitat use by small cetaceans within shelf-seas is temporally variable, species specific and likely driven by complex bottom-up processes. As such, the effective conservation management of shelf-seas requires that we understand the dynamic complexities of these systems and the species that inhabit them. In particular, we emphasise the need for a good understanding of the natural drivers of habitat use by marine megafauna before the potential impacts of anthropogenically induced disturbances, such as those associated with the construction, maintenance and operation of MREIs, can be assessed.

Background. Citizen science is increasingly popular and has the potential to collect extensive datasets at lower costs than traditional surveys. Ferries have been used to collect data on cetacean populations for decades, providing long-term time series allowing for monitoring of cetacean populations. One cetacean species of concern is the common dolphin, which have been found stranded around the north-east Atlantic in recent years, with high numbers on French coasts being attributed to fisheries bycatch. We estimate common dolphin densities in north-east Atlantic and investigate the power of citizen science data to identify changes in marine mammal densities and areas of importance. Materials & Methods. Data were collected by citizen scientists on ferries between April and October in 2006 - 2017. Common dolphin sightings data from two ferry routes in the Bay of Biscay (n= 569), Celtic Sea (n= 260), and English Channel (n= 75) were used to estimate detection probabilities with detection functions. Density Surface Models estimated density across ferry routes, accounting for the influence of environmental (chlorophyll a, sea surface temperature, depth, and slope), spatial (latitude and longitude) and temporal terms (year and Julian day). Results. Overall detection probability was highest in the English Channel (0.384) and Bay of Biscay (0.348), and lowest in the Celtic Sea (0.158). Common dolphins were estimated to occur in higher densities in the Celtic Sea (0.400 per km) and the Bay of Biscay (0.319 per km), with low densities in the English Channel (0.025 per km). Densities in the Celtic Sea have been relatively stable on the ferry route since 2006 with a slight decrease in 2017. Densities peaked in the Bay of Biscay in 2013 with lower numbers since. The general trend in the English Channel is for increasing densities of common dolphins over time since 2009. Discussion. This study highlights the effectiveness of citizen science data to investigate the distribution and density of cetaceans. The densities and temporal changes shown by this study are representative of those from wider-ranging robust estimates. We highlight the ability of citizen science to collect data over extensive periods of time which complements traditional surveys. Such long-term data are important to identify changes within a population; however, citizen science data may, in some situations, present challenges. We provide recommendations to ensure high-quality data which can be used to inform management and conservation of cetacean populations.

For much of the twentieth century, many or most scholarly journals in biodiversity, ecology, biogeography and conservation were owned and published by scientific societies and made available to the broader scientific community at relatively low cost. However, the past several decades have seen a dominant process of commercialization of the publication process in this field, either by commercial publishing firms taking over publication of journals owned by a society (e.g., Evolution), or by commercial publishers starting new journals to fill “niches” that were until then empty in the scholarly publishing ecosystem. Diversity and Distributions is an example of this latter category of journals, having been created by Wiley Publishers in 1993 (as Biodiversity Letters). Being a journal owned by a commercial publisher, it has always been accessed by readers via subscription, but the publication process has been free to potential authors whose work has passed peer review.

The wave energy industry is an emerging sector and a new user of maritime space that has potential to contribute significantly to the EU renewable energy goals. International and national regulatory frameworks necessitate Environmental Impact Assessments (EIA) that provide important data to inform development consent decisions. Here we have evaluated experience related to the assessment programmes at EU wave energy test centres combined with knowledge gained from EIA produced for other similar renewable energy developments. From this we have identified key receptors of concern, as well as the type and magnitude of impacts which may be expected. The key environmental receptors of concern for wave energy EIA include the physical environment (e.g. morphology, waves and current) and flora and fauna 1 as represented by marine mammals, seabirds, benthos, fish and shellfish. From a review of the EIAs performed at wave energy test centres, we identified several lessons regarding the wave energy EIA process. There is clear evidence that the receptors of primary interest are dependent on factors such as the local environmental characteristics, the presence/absence of protected species and the regulatory authority under which the EIA is performed. Furthermore, it is recommended that concerns relating to cumulative impacts, from an expanding level of wave energy development taking place in a background of growing utilisation of the marine environment, which are largely unknown at this early stage of the industry may be comprehensively addressed at the national level as part of a Strategic Environmental Assessment (EIA) and/or in Maritime Spatial Planning (MSP) and that it should be regularly reassessed.

Background Citizen science is increasingly popular and has the potential to collect extensive datasets at lower costs than traditional surveys conducted by professional scientists. Ferries have been used to collect data on cetacean populations for decades, providing long-term time series for monitoring of cetacean populations. One cetacean species of concern is the common dolphin, which has been found stranded around the north-east Atlantic in recent years, with high numbers on French coasts being attributed to fisheries bycatch. We estimate common dolphin densities in the north-east Atlantic and investigate the ability of citizen science data to identify changes in marine mammal densities and areas of importance. Materials and Methods Data were collected by citizen scientists on ferries between April and October in 2006–2017. Common dolphin sightings data from two ferry routes across three regions, Bay of Biscay (n = 569); south-west United Kingdom to the Isles of Scilly in the Celtic Sea (n = 260); and English Channel (n = 75), were used to estimate density across ferry routes. Two-stage Density Surface Models accounted for imperfect detection, and tested the influence of environmental (chlorophyll a, sea surface temperature, depth, and slope), spatial (latitude and longitude) and temporal terms (year and Julian day) on occurrence. Results Overall detection probability was highest in the areas sampled within the English Channel (0.384) and Bay of Biscay (0.348), and lowest on the Scilly’s route (0.158). Common dolphins were estimated to occur in higher densities on the Scilly’s route (0.400 per km2) and the Bay of Biscay (0.319 per km2), with low densities in the English Channel (0.025 per km2). Densities on the Scilly’s route appear to have been relatively stable since 2006 with a slight decrease in 2017. Densities peaked in the Bay of Biscay in 2013 with lower numbers since. Densities in the English Channel appear to have increased over time since 2009. Discussion This study highlights the effectiveness of citizen science data to investigate the distribution and density of cetaceans. The densities and temporal changes shown by this study are representative of those from wider-ranging robust estimates. We highlight the ability of citizen science to collect data over extensive periods of time which complements dedicated, designed surveys. Such long-term data are important to identify changes within a population; however, citizen science data may, in some situations, present challenges. We provide recommendations to ensure high-quality data which can be used to inform management and conservation of cetacean populations.

International audience; Predictive models are central to many scientific disciplines and vital for informing management in a rapidly changing world. However, limited understanding of the accuracy and precision of models transferred to novel conditions (their 'trans-ferability') undermines confidence in their predictions. Here, 50 experts identified priority knowledge gaps which, if filled, will most improve model transfers. These are summarized into six technical and six fundamental challenges, which underlie the combined need to intensify research on the determinants of ecological predictability, including species traits and data quality, and develop best practices for transferring models. Of high importance is the identification of a widely applicable set of transferability metrics, with appropriate tools to quantify the sources and impacts of prediction uncertainty under novel conditions. Predicting the Unknown Predictions facilitate the formulation of quantitative, testable hypotheses that can be refined and validated empirically [1]. Predictive models have thus become ubiquitous in numerous scientific disciplines, including ecology [2], where they provide means for mapping species distributions, explaining population trends, or quantifying the risks of biological invasions and disease outbreaks (e.g., [3,4]). The practical value of predictive models in supporting policy and decision making has therefore grown rapidly (Box 1) [5]. With that has come an increasing desire to predict (see Glossary) the state of ecological features (e.g., species, habitats) and our likely impacts upon them [5], prompting a shift from explanatory models to anticipatory predictions [2]. However, in many situations, severe data deficiencies preclude the development of specific models, and the collection of new data can be prohibitively costly or simply impossible [6]. It is in this context that interest in transferable models (i.e., those that can be legitimately projected beyond the spatial and temporal bounds of their underlying data [7]) has grown. Transferred models must balance the tradeoff between estimation and prediction bias and variance (homogenization versus nontransferability, sensu [8]). Ultimately, models that can Highlights Models transferred to novel conditions could provide predictions in data-poor scenarios, contributing to more informed management decisions.

Tags, and their deployments, were funded by the Welsh Assembly Government (Welsh colonies; project no. JER3688), Marine Scotland (Stroma and Muckle Green Holm; project no. CR/2009/48), the Natural Environment Research Council (NERC) (Isle of May) and SMRU Instrumentation (Isle of May). MIDC studentship is co-funded by Plymouth University School of Biological & Marine Sciences and NERC. DJFR, CJB & DT are supported by National Capability funding from NERC to SMRU (grant no. SMRU1001). Young animals must learn to forage effectively to survive the transition from parental provisioning to independent feeding. Rapid development of successful foraging strategies is particularly important for capital breeders that do not receive parental guidance after weaning. The intrinsic and extrinsic drivers of variation in ontogeny of foraging are poorly understood for many species. Grey seals (Halichoerus grypus) are typical capital breeders; pups are abandoned on the natal site after a brief suckling phase, and must develop foraging skills without external input. We collected location and dive data from recently-weaned grey seal pups from two regions of the United Kingdom (the North Sea and the Celtic and Irish Seas) using animal-borne telemetry devices during their first months of independence at sea. Dive duration, depth, bottom time, and benthic diving increased over the first 40 days. The shape and magnitude of changes differed between regions. Females consistently had longer bottom times, and in the Celtic and Irish Seas they used shallower water than males. Regional sex differences suggest that extrinsic factors, such as water depth, contribute to behavioural sexual segregation. We recommend that conservation strategies consider movements of young naïve animals in addition to those of adults to account for developmental behavioural changes. Publisher PDF

Young animals must learn to forage effectively to survive the transition from parental provisioning to independent feeding. Rapid development of successful foraging strategies is particularly important for capital breeders that do not receive parental guidance after weaning. The intrinsic and extrinsic drivers of variation in ontogeny of foraging are poorly understood for many species. Grey seals (Halichoerus grypus) are typical capital breeders; pups are abandoned on the natal site after a brief suckling phase, and must develop foraging skills without external input. We collected location and dive data from recently-weaned grey seal pups from two regions of the United Kingdom (the North Sea and the Celtic and Irish Seas) using animal-borne telemetry devices during their first months of independence at sea. Dive duration, depth, bottom time, and benthic diving increased over the first 40 days. The shape and magnitude of changes differed between regions. Females consistently had longer bottom times, and in the Celtic and Irish Seas they used shallower water than males. Regional sex differences suggest that extrinsic factors, such as water depth, contribute to behavioural sexual segregation. We recommend that conservation strategies consider movements of young naïve animals in addition to those of adults to account for developmental behavioural changes.

As we begin to manage our oceans in much more spatial detail we must understand a great deal more about oceanographic habitat preferences of marine mobile top predators. In this unique field study we test a hypothesis on the mechanisms defining mobile predator foraging habitat characteristics by comparing temporally and spatially detailed bio-physical oceanographic data from contrasting topographical locations. We contrast the foraging locations of two very different seabird species, gannets and storm petrels, by repeatedly sampling a bank and a nearby flat area over daily tidal cycles during spring and neap tides. The results suggest that storm petrels are linked to foraging in specific locations where internal waves are produced, which is mainly on banks. These locations can also include the presence of high biomass of chlorophyll. In contrast, the location where more gannets are foraging is significantly influenced by temporal variables with higher densities of foraging birds much more likely during the neap tide than times of spring tide. The foraging times of both species was influenced by differences between the vertical layers of the water column above and below the thermocline; via either vertical shear of horizontal currents or absolute differences in speed between layers. Higher densities of foraging gannets were significantly more likely to be found at ebb tides in both bank and flat regions however over the bank, the density of foraging gannets was higher when the differences in speed between the layers were at a maximum. Both gannets and storm petrels appear to be more likely to forage when wind direction is opposed to tidal direction. This detailed understanding links foraging behaviour to predictable spatial and temporal bio-physical vertical characteristics and thus can be immediately used to explain variance and increase certainty in past abundance and distributional surveys. These results also illuminate the types of variables that should be considered when assessing potential changes to the distribution and characteristics of habitats from increased anthropogenic disturbances such as large scale offshore wind, wave and tidal renewable deployments.

The availability of fish to top mobile predators such as seabirds and fishermen and to fisheries surveys depends on their schooling behaviour. In temperate coastal ecosystems, tide is the main driver of ecosystem dynamics. To investigate the mechanisms linking fish schooling behaviour to bio-physical oceanography in a tidal ecosystem, we combined novel fine scale survey data of fish behaviour using fisheries acoustics with co-located oceanographic data from contrasting topographical locations. The schooling behaviour of pelagic fish was examined in relation to bio-physical oceanography by carrying out repeated surveys of the same locations over the daily tidal cycle at spring and neap tides over a small bank (Jones Bank) and a nearby flat region. Different fish species were found over the bank and in the flat region 14 km to the south-east, and fish schooling behaviour varied similarly. Results showed that the bank played an important role in influencing fish distribution and behaviour, with shallow pelagic schools concentrated over the bank and closer to the surface at times of high internal wave activity. Behaviour of schools identified close to the bottom was partially influenced by oceanographic variables that drove internal waves over the bank slopes, but were most strongly influenced by the spring–neap cycle. These deep schools were larger, closer to the bottom and less dense at neap tides. Identifying some the of the bio-physical variables which drive changes in fish distribution and behaviour will help quantify fish catchability and assist in the design and interpretation of better targeted surveys for top predators and prey fish species.

This work was funded under a NERC studentship to the lead author with CASE funding from Scottish Natural Heritage. The Hebridean Whale and Dolphin Trust were funded by the Heritage Lottery Fund, Scottish Natural Heritage, Argyll and the Islands Enterprise, WWF and the Earthwatch Institute. The west coast of Scotland is comprised of complex coastlines and topography, and a range of physical processes influence its coastal marine environment. The region is host to one of the highest densities of harbour porpoise Phocoena phocoena in Europe. The aim of this study was to identify habitat preferences driving the distribution of harbour porpoise, to gain a better understanding of the spatial distribution of the species in the region, as well as to assess the consistency of such patterns across time and space. Visual and acoustic line-transect surveys were conducted between 2003 and 2010. Generalised Additive Models (GAMs) with Generalised Estimating Equations (GEEs) were used to robustly determine relationships between the relative density of harbour porpoises and temporally and spatially variable oceanographic covariates. Predictive models showed that depth, slope, spring tidal range and distance to land were consistently important in explaining porpoise distribution. Consistent preferences for water depths between 50 and 150 m and highly sloped regions were observed across the temporal models. Predicted distributions revealed a consistent inshore presence for the species throughout the west coast of Scotland and confirmed that predictable oceanographic features could help inform the establishment of Special Areas of Conservation (SACs) for the species. Publisher PDF

This study was supported by the EU (via PERSEUS grant FP7-OCEAN-2011-287600 and MyOcean SPA.2011.1.5-01 grant 283367). In addition, C.B. Embling was supported by the 2014 SoMSE Plymouth University Small (grant number 2014-SoMSE) research grant. D.J.F. Russell and D. Thompson were supported by the National Capability funding from the Natural Environment Research Council to the Sea Mammal Research Unit (grant no. SMRU1001). C. Vincent was supported by the Parc naturel marin d'Iroise and Région Poitou – Charentes (grant number 782423). Shipping noise is a threat to marine wildlife. Grey seals are benthic foragers, and thus experience acoustic noise throughout the water column, which makes them a good model species for a case study of the potential impacts of shipping noise. We used ship track data from the Celtic Sea, seal track data and a coupled ocean-acoustic modelling system to assess the noise exposure of grey seals along their tracks. It was found that the animals experience step changes in sound levels up to ~ 20 dB at a frequency of 125 Hz, and ~ 10 dB on average over 10–1000 Hz when they dive through the thermocline, particularly during summer. Our results showed large seasonal differences in the noise level experienced by the seals. These results reveal the actual noise exposure by the animals and could help in marine spatial planning. Postprint

1. Spatial management of marine ecosystems requires detailed knowledge of spatio-temporal mechanisms linking physical and biological processes. Tidal currents, the main driver of ecosystem dynamics in temperate coastal ecosystems, influence predator foraging ecology by affecting prey distribution and ecology. The mechanistic links between tidal currents and how they influence predator–prey behaviour and interactions at a fine scale are poorly understood. 2. Studies of fine-scale changes in oceanography, prey and predator behaviour with tidal currents require repeated surveys of the same location over brief time-scales. Such data are highly temporally and spatially autocorrelated and require appropriate analytical tools. 3. We used functional data analysis (FDA), specifically functional principal component analysis (FPCA), to analyse repeated, fine-scale, survey data collected in the North Sea. FPCA was used to explore the relationship between the behaviour of an important North Sea prey species (sandeel Ammodytes spp.) and a vulnerable surface-foraging predator (black-legged kittiwake Rissa tridactyla) with fine-scale tidally driven changes in bio-physical characteristics (temperature stratification and maximumsubsurface chlorophyll concentration). 4. The FPCA indicated that sandeels were aggregated close to the surface at maximum ebb (ME) currents. Surface-feeding kittiwakes were also found in highest numbers during ME in locations of both high subsurface chlorophyll concentration and shallow sandeel aggregations. We suggest that the combination of a well-stratified water column with the movement of tidal currents over uneven topography results in surface aggregations of sandeels which kittiwakes exploit. 5. Synthesis and applications. Functional Data Analysis provides a useful tool for examining spatio-temporal patterns in natural ecosystems. In combination with fine-scale repeated survey design, we identified the importance of tide in driving prey behaviour and hence predator foraging behaviour. This has implications both for critical marine habitat identification for Marine Protected Area selection and for fisheries stock assessments. We therefore recommend that tidal aspects should be taken into account when designing marine surveys in temperate coastal ecosystems both to ensure the best identification of critical marine habitat and to improve the accuracy of fish stock assessments.

The harbour porpoise is a highly mobile species and thus represents a considerable challenge in the context of using marine protected areas (MPAs) for conservation. The shelf waters off the west coast of Scotland have been identified as an area of year-round presence, high density in comparison to surrounding areas, and a high young to adult ratio in summer and are thus a suitable area for exploring the location of possible special areas of conservations (SACs) under the EU Habitats Directive. We carried out dedicated surveys over three summers in the southern Inner Hebrides and used generalised additive models (GAMs) to predict areas of high relative density for harbour porpoises for each year. After compensating for survey effects, static bathymetric and persistent hydrographic variables were used in a step-wise model selection procedure. In all years harbour porpoise distribution was best explained by maximum tidal current, with higher densities predicted in areas of low current, and the same high density areas predicted year-on-year. Perimeter-to-area ratio was used to identify which areas should be considered as a basis for designating SACs for harbour porpoise in this area, to form part of the Natura 2000 network. The method used here combines spatial modelling and perimeter-to-area ratio for selecting protected areas, a methodology which is suitable for the protection of other animal species.

Growing concerns about climate change and energy security have fueled a rapid increase in the development of marine renewable energy installations (MREIs). The potential ecological consequences of increased use of these devices emphasizes the need for high quality environmental impact assessment (EIA). We demonstrate that these processes are hampered severely, primarily because ambiguities in the legislation and lack of clear implementation guidance are such that they do not ensure robust assessment of the significance of impacts and cumulative effects. We highlight why the regulatory framework leads to conceptual ambiguities and propose changes which, for the most part, do not require major adjustments to standard practice. We emphasize the importance of determining the degree of confidence in impacts to permit the likelihood as well as magnitude of impacts to be quantified and propose ways in which assessment of population-level impacts could be incorporated into the EIA process. Overall, however, we argue that, instead of trying to ascertain which particular developments are responsible for tipping an already heavily degraded marine environment into an undesirable state, emphasis should be placed on better strategic assessment. Publisher PDF

The oceanographic drivers of marine vertebrate habitat use are poorly understood yet fundamental to our knowledge of marine ecosystem functioning. Here, we use composite front mapping and high-resolution GPS tracking to determine the significance of mesoscale oceanographic fronts as physical drivers of foraging habitat selection in northern gannets Morus bassanus . We tracked 66 breeding gannets from a Celtic Sea colony over 2 years and used residence time to identify area-restricted search (ARS) behaviour. Composite front maps identified thermal and chlorophyll- a mesoscale fronts at two different temporal scales—(i) contemporaneous fronts and (ii) seasonally persistent frontal zones. Using generalized additive models (GAMs), with generalized estimating equations (GEE-GAMs) to account for serial autocorrelation in tracking data, we found that gannets do not adjust their behaviour in response to contemporaneous fronts. However, ARS was more likely to occur within spatially predictable, seasonally persistent frontal zones (GAMs). Our results provide proof of concept that composite front mapping is a useful tool for studying the influence of oceanographic features on animal movements. Moreover, we highlight that frontal persistence is a crucial element of the formation of pelagic foraging hotspots for mobile marine vertebrates.

The coastal Runnelstone Reef, off southwest Cornwall (UK), is characterised by complex topography and strong tidal flows and is a known high-density site for harbour porpoise (Phocoena phocoena); a European protected species. Using a multidisciplinary dataset including: porpoise sightings from a multi-year land-based survey, Acoustic Doppler Current Profiling (ADCP), vertical profiling of water properties and high-resolution bathymetry; we investigate how interactions between tidal flow and topography drive the fine-scale porpoise spatio-temporal distribution at the site. Porpoise sightings were distributed non-uniformly within the survey area with highest sighting density recorded in areas with steep slopes and moderate depths. Greater numbers of sightings were recorded during strong westward (ebbing) tidal flows compared to strong eastward (flooding) flows and slack water periods. ADCP and Conductivity Temperature Depth (CTD) data identified fine-scale hydrodynamic features, associated with cross-reef tidal flows in the sections of the survey area with the highest recorded densities of porpoises. We observed layered, vertically sheared flows that were susceptible to the generation of turbulence by shear instability. Additionally, the intense, oscillatory near surface currents led to hydraulically controlled flow that transitioned from subcritical to supercritical conditions; indicating that highly turbulent and energetic hydraulic jumps were generated along the eastern and western slopes of the reef. The depression and release of isopycnals in the lee of the reef during cross-reef flows revealed that the flow released lee waves during upslope currents at specific phases of the tidal cycle when the highest sighting rates were recorded. The results of this unique, fine-scale field study provide new insights into specific hydrodynamic features, produced through tidal forcing, that may be important for creating predictable foraging opportunities for porpoises at a local scale. Information on the functional mechanisms linking porpoise distribution to static and dynamic physical habitat variables is extremely valuable to the monitoring and management of the species within the context of European conservation policies and marine renewable energy infrastructure development.

The climatic changes of the glacial cycles are thought to have been a major driver of population declines and species extinctions. However, studies to date have focused on terrestrial fauna and there is little understanding of how marine species responded to past climate change. Here we show that a true Arctic species, the bowhead whale (Balaena mysticetus), shifted its range and tracked its core suitable habitat northwards during the rapid climate change of the Pleistocene-Holocene transition. Late Pleistocene lineages survived into the Holocene and effective female population size increased rapidly, concurrent with a threefold increase in core suitable habitat. This study highlights that responses to climate change are likely to be species specific and difficult to predict. We estimate that the core suitable habitat of bowhead whales will be almost halved by the end of this century, potentially influencing future population dynamics.

The first recordings from free-ranging Gervais' beaked whale (Mesoplodon europaeus) are presented. Nine Gervais' beaked whales were observed visually for over 6 h. Clicks were only detected over a 15 min period during the encounter, which coincided with an 88 min period during which no whales were observed at the surface. Click lengths were typically around 200 microS and their dominant energy was in the frequency range 30-50 kHz. While these clicks were broadly similar to those of Cuvier's and Blainville's beaked whales, the Gervais' beaked whale clicks were at a slightly higher frequency than those of the other species.

Oceanic fronts are key habitats for a diverse range of marine predators, yet how they influence fine-scale foraging behaviour is poorly understood. Here, we investigated the dive behaviour of northern gannets Morus bassanus in relation to shelf-sea fronts. We GPS (global positioning system) tracked 53 breeding birds and examined the relationship between 1901 foraging dives (from time-depth recorders) and thermal fronts (identified via Earth Observation composite front mapping) in the Celtic Sea, Northeast Atlantic. We (i) used a habitat-use availability analysis to determine whether gannets preferentially dived at fronts, and (ii) compared dive characteristics in relation to fronts to investigate the functional significance of these oceanographic features. We found that relationships between gannet dive probabilities and fronts varied by frontal metric and sex. While both sexes were more likely to dive in the presence of seasonally persistent fronts, links to more ephemeral features were less clear. Here, males were positively correlated with distance to front and cross-front gradient strength, with the reverse for females. Both sexes performed two dive strategies: shallow V-shaped plunge dives with little or no active swim phase (92% of dives) and deeper U-shaped dives with an active pursuit phase of at least 3 s (8% of dives). When foraging around fronts, gannets were half as likely to engage in U-shaped dives compared with V-shaped dives, independent of sex. Moreover, V-shaped dive durations were significantly shortened around fronts. These behavioural responses support the assertion that fronts are important foraging habitats for marine predators, and suggest a possible mechanistic link between the two in terms of dive behaviour. This research also emphasizes the importance of cross-disciplinary research when attempting to understand marine ecosystems.