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Funding: This work was funded by the Office of Naval Research under award: N000141912464. 1. The assessment of behavioural disturbance in cetacean species (e.g., resulting from exposure to anthropogenic sources such as military sonar, seismic surveys, or pile driving) is important for effective conservation and management. Disturbance effects can be informed by Behavioural Response Studies (BRSs), involving either controlled exposure experiments (CEEs) where noise exposure conditions are presented deliberately to meet experimental objectives or in opportunistic contexts where ongoing activities are monitored in a strategic manner. In either context, animal-borne sensors or in-situ observations can provide information on individual exposure and disturbance responses. 2. The past 15 years of research have greatly expanded our understanding of behavioural responses to noise, including hundreds of experiments in nearly a dozen cetacean species. Many papers note limited sample sizes, required knowledge of baseline behaviour prior to exposure and the importance of contextual factors modulating behavioural responses, all of which in combination can lead to sampling biases, even for well-designed research programs. 3. It is critical to understand these biases to robustly identify responses. This ensures outcomes of BRSs help inform predictions of how anthropogenic disturbance impacts individuals and populations. Our approach leverages concepts from the animal behaviour literature focused on helping to avoid sampling bias by considering what shapes an animal's response. These factors include social, experience, genetic and natural changes in responsiveness. 4. We developed and applied a modified version of this framework to synthesize current knowledge on cetacean response in the context of effects observed across marine and terrestrial taxa. This new 'Sampling, Exposure, Receptor' framework (SERF) identifies 43 modulating factors, highlights potential biases, and assesses how these vary across selected focal species. 5. In contrast to studies that identified variation in 'Exposure' factors as a key concern, our analysis indicated that factors relating to 'Sampling' (e.g., deploying tags on less evasive individuals, which biases selection of subjects), and "Receptor" (e.g., health status or coping style) have the greatest potential for weakening the desired broad representativeness of BRSs. Our assessment also highlights how potential biases could be addressed with existing datasets or future developments. Publisher PDF

Bioenergetics is the study of how animals achieve energetic balance. Energetic balance results from the energetic expenditure of an individual and the energy they extract from their environment. Ingested energy depends on several extrinsic (e.g prey species, nutritional value and composition, prey density and availability) and intrinsic factors (e.g. foraging effort, success at catching prey, digestive processes and associated energy losses, and digestive capacity). While the focus in bioenergetic modelling is often on the energetic costs an animal incurs, the robust estimation of an individual’s energy intake is equally critical for producing meaningful predictions. Here, we review the components and processes that affect energy intake from ingested gross energy to biologically useful net energy (NE). The current state of knowledge of each parameter is reviewed, shedding light on research gaps to advance this field. The review highlighted that the foraging behaviour of many marine mammals is relatively well studied via biologging tags, with estimates of success rate typically assumed for most species. However, actual prey capture success rates are often only assumed, although we note studies that provide approaches for its estimation using current techniques. A comprehensive collation of the nutritional content of marine mammal prey species revealed a robust foundation from which prey quality (comprising prey species, size and energy density) can be assessed, though data remain unavailable for many prey species. Empirical information on various energy losses following ingestion of prey was unbalanced among marine mammal species, with considerably more literature available for pinnipeds. An increased understanding and accurate estimate of each of the components that comprise a species NE intake are an integral part of bioenergetics. Such models provide a key tool to investigate the effects of disturbance on marine mammals at an individual and population level and to support effective conservation and management.

We wish to thank Fundación Biodiversidad for supporting COLCA project and all the ecovolunteers involved in the fieldwork of the Balearic Sperm Whale project. We are grateful to OceanCare (Switzerland) for their continuous support of the sperm whale research and conservation activities of the PCRI since 2008 and to Prof. Kostas Kostarelos and the University of Manchester for organizing the crowd-funding project “Nanowhales” to cofund the research expedition of 2019 along the Hellenic Trench. This paper represents HIMB and SOEST contribution nos. 1903 and 11568, respectively. Body mass is a fundamental feature of animal physiology. Although sperm whales (Physeter macrocephalus) are the largest toothed predators on earth, body mass is seldom included in studies of their ecophysiology and bioenergetics due to the inherent difficulties of obtaining direct measurements. We used UAV‐photogrammetry to estimate the weight of free‐ranging sperm whales. Aerial photographs (23 calves, 11 juveniles, 55 nonmother adults, 13 mothers) were collected in the Eastern Caribbean and Mediterranean Sea during 2017–2020. Body length, widths, and heights (dorso‐ventral distance at 5% increments) were measured from dorsal and lateral photographs, while body volume was calculated using an elliptical model. Volume varied noticeably (12.01 ± 4.79 m3) in larger animals (>8 m), indicating fluctuations in body condition of adults and mothers. Volume was converted to mass, using tissue‐density estimates from catch data, animal‐borne tags, and body‐tissue composition. Average total body density ranged from 834 to 1,003 kg/m3, while the weight predictions matched with existing measurements and weight‐length relationships. Our body‐mass models can be used to study sperm whale bioenergetics, including inter‐ and intraseasonal variations in body condition, somatic growth, metabolic rates, and cost of reproduction. Publisher PDF

Funding: J.H.D. was funded by the Irish Research Council Enterprise Partnership Scheme, supported by the Petroleum Infrastructure Program. Field work on Little Saltee in 2018 and 2019 and S.d.G. were funded by the BlueFish project, funded by the European Regional Development fund through the Ireland Wales Cooperation Programme 2014−2020. Fieldwork on Eynhallow and St. Kilda was supported by Orkney Islands Council, the University of Aberdeen, the National Trust for Scotland and Talisman Energy (UK) Ltd. E.W.J.E. was funded by a Marine Alliance for Science and Technology for Scotland and University of Aberdeen studentship. Fieldwork elsewhere was funded by the EU Atlantic area INTERREG program via the Future of the Atlantic Marine Environment (FAME) project and by the RSPB, JNCC, Fair Isle Bird Observatory Trust and Marine Scotland, through the Seabird Tracking And Research (STAR) project. G.E.A. was funded by the MarPAMM project supported by the EU INTERREG VA Programme, managed by the Special EU Programmes Body (SEUPB). Habitat-use and distribution models are essential tools of conservation biology. For wide-ranging species, such models may be challenged by the expanse, remoteness and variability of their habitat, these challenges often being compounded by the species' mobility. In marine environments, direct observations and sampling are usually impractical over broad regions, and instead remotely sensed proxies of prey availability are often used to link species abundance or foraging behaviour to areas that are expected to provide food consistently. One source of food consumed by many marine top predators is fisheries waste, but habitat-use models rarely account for this interaction. We assessed the utility of commercial fishing effort as a covariate in foraging habitat models for northern fulmars Fulmarus glacialis, a species known to exploit fisheries waste, during their summer breeding season. First, we investigated the prevalence of fulmar-vessel interactions using concurrently tracked fulmars and fishing vessels. We infer that over half of our study individuals associate with fishing vessels while foraging, mostly with trawl-type vessels. We then used hidden Markov models to explain the spatio-temporal distribution of putative foraging behaviour as a function of a range of covariates. Persistent commercial fishing effort was a significant predictor of foraging behaviour, and was more important than commonly used environmental covariates retained in the model. This study demonstrates the effect of commercial fisheries on the foraging distribution and behaviour of a marine top predator, and supports the idea that, in some systems, incorporating human activities into distribution studies can improve model fit substantially. Publisher PDF

Funding: Office of Naval Research - N000142012697, N000142112096; Strategic Environmental Research and Development Program - RC20-1097, RC20-7188, RC21-3091. Assessing cumulative effects of human activities on ecosystems is required by many jurisdictions, but current science cannot meet regulatory demands. Regulations define them as effect(s) of one human action combined with other actions. Here we argue for an approach that evaluates the cumulative risk of multiple stressors for protected wildlife populations within their ecosystems. Monitoring effects of each stressor is necessary but not sufficient to estimate how multiple stressors interact to affect wildlife populations. Examining the mechanistic pathways, from cellular to ecological, by which stressors affect individuals can help prioritize stressors and interpret how they interact. Our approach uses health indicators to accumulate the effects of stressors on individuals and to estimate changes in vital rates, driving population status. We advocate using methods well-established in human health and integrating them into ecosystem-based management to protect the health of commercially and culturally important wildlife populations and to protect against risk of extinction for threatened species. Our approach will improve abilities to conserve and manage ecosystems but will also demand significant increases in research and monitoring effort. We advocate for increased investment proportional to the economic scale of human activities in the Anthropocene and their pervasive effects on ecology and biodiversity. Publisher PDF

This work was funded by the Marine Mammal Commission (MMC19-173). The Office of Naval Research funded the bioenergetic workshop (N000142012392) that provided support for this work. Bioenergetic approaches are increasingly used to understand how marine mammal populations could be affected by a changing and disturbed aquatic environment. There remain considerable gaps in our knowledge of marine mammal bioenergetics, which hinder the application of bioenergetic studies to inform policy decisions. We conducted a priority-setting exercise to identify high-priority unanswered questions in marine mammal bioenergetics, with an emphasis on questions relevant to conservation and management. Electronic communication and a virtual workshop were used to solicit and collate potential research questions from the marine mammal bioenergetic community. From a final list of 39 questions, 11 were identified as ‘key’ questions because they received votes from at least 50% of survey participants. Key questions included those related to energy intake (prey landscapes, exposure to human activities) and expenditure (field metabolic rate, exposure to human activities, lactation, time-activity budgets), energy allocation priorities, metrics of body condition and relationships with survival and reproductive success and extrapolation of data from one species to another. Existing tools to address key questions include labelled water, animal-borne sensors, mark-resight data from long-term research programs, environmental DNA and unmanned vehicles. Further validation of existing approaches and development of new methodologies are needed to comprehensively address some key questions, particularly for cetaceans. The identification of these key questions can provide a guiding framework to set research priorities, which ultimately may yield more accurate information to inform policies and better conserve marine mammal populations. Publisher PDF

Funding: This work was supported by the National Geographic Society [grant number EC-53116R-18]. Sexual segregation is common among marine mammals, leading to intraspecific differences in diet, diving behaviour, home range size and even latitudinal distribution and migratory patterns. Sperm whales Physeter macrocephalus present one of the most extreme examples of sexual dimorphism both in size and social structure, with males and females segregating at different latitudes across most of their range, but the underlying ecological drivers remain unclear. Studying fine-scale dietary and habitat differences where the sexes occur in sympatry could therefore provide insights into the mechanisms underpinning their large-scale segregation. In this study, we analysed the carbon and nitrogen stable isotope values in the skin of males and females from an isolated, endangered population inhabiting the Mediterranean Sea, sampled in a region where the sexes occur and feed regularly in the summer months but show subtle differences in habitat preference. We found marked differences in both carbon and nitrogen isotopic values between the sexes, indicating that they could be targeting prey items in different trophic levels and habitats. Combined with the evidence from habitat modelling studies, our results suggest that female and male sperm whales segregate even in the latitudinally restricted Mediterranean population, at a much smaller scale. This sympatric, fine-scale sexual segregation suggests that reduction of competition may have been a key factor in the evolution of the social structure and large-scale latitudinal segregation of this species. Postprint

Predicting the distribution and behaviour of animals is a fundamental objective in ecology and a cornerstone of conservation biology. Modelling the distribution of ocean-faring species like seabirds remains a significant challenge due to ocean dynamics, colony-specific effects and the vast ranges seabirds can cover. We used a spatial and behavioural approach to model the distribution of the Manx shearwater Puffinus puffinus, a pelagic, central-place forager that can cover great distances while foraging. GPS data from birds tagged in 2 colonies over 3 yr were modelled with a range of environmental predictors of marine productivity. For both colonies, transitions to foraging behaviour correlated with chlorophyll a, and the distribution of foraging behaviour was also associated with areas of high chlorophyll a concentration in coastal but not offshore areas for one colony. Furthermore, there was evidence for colony differences in habitat use, prevalence of nocturnal foraging, and for some competitive exclusion on foraging grounds, even though the colonies were 170 km apart. Despite the extensive dataset, our models had modest predictive power, which we suggest can probably only be improved by including biotic interactions, including more direct measures of food resource distribution. Our results highlight the importance of including spatial complexity and data from multiple sites when predicting the distribution of wide-ranging predators, because patterns of distribution and habitat use likely differ across the range of a population.

This work was supported by the United States Office of Naval Research (awards N00014-17-1-2757 and N00014-19-1-2479). Lipid-store body condition is fundamental to how animals cope with environmental fluctuations, including anthropogenic change. As it provides an energetic buffer, body condition is expected to influence risk-taking strategies, with both positive and negative relationships between body condition and risk-taking posited in the literature. Individuals in good condition may take more risks due to state-dependent safety (‘ability-based’ explanation), or alternatively fewer risks due to asset protection and reduced need to undertake risky foraging (‘needs-based’ explanation). Such state-dependent responses could drive non-linear impacts of anthropogenic activities through feedback between body condition and behavioural disturbance. Here, we present a simple bioenergetic model that explicitly incorporates hypothetical body condition-dependent response strategies for a cetacean, the sperm whale. The model considered the consequences of state-dependent foraging cessation and availability of wax ester (WE) lipids for calf provisioning and female survival. We found strikingly different consequences of disturbance depending on strategy and WE availability scenarios. Compared with the null strategy, where responses to disturbance were independent of body condition, the needs-based strategy mitigated predicted reductions in provisioning by 10%–13%, while the ability-based strategy exaggerated reductions by 63%–113%. Lower WE availability resulted in more extreme outcomes because energy stores were smaller relative to the daily energy balance. In the 0% availability scenario, while the needs-based strategy reduced deaths by 100%, the ability-based strategy increased them by 335% relative to null and by 56% relative to the same strategy under the 5%–6.7% WE availability scenario. These results highlight that state-dependent disturbance responses and energy store availability could substantially impact the population consequences of disturbance. Our ability to set appropriate precautionary disturbance thresholds therefore requires empirical tests of ability- vs needs-based response modification as a function of body condition and a clearer understanding of energy store availability. Publisher PDF

Bioenergetic models describe the processes through which animals acquire energy from resources in the environment and allocate it to different life history functions. They capture some of the fundamental mechanisms regulating individuals, populations and ecosystems and have thus been used in a wide variety of theoretical and applied contexts. Here, I review the development of bioenergetic models for marine mammals and their application to management and conservation. For these long-lived, wide-ranging species, bioenergetic approaches were initially used to assess the energy requirements and prey consumption of individuals and populations. Increasingly, models are developed to describe the dynamics of energy intake and allocation and predict how resulting body reserves, vital rates and population dynamics might change as external conditions vary. The building blocks required to develop such models include estimates of intake rate, maintenance costs, growth patterns, energy storage and the dynamics of gestation and lactation, as well as rules for prioritizing allocation. I describe how these components have been parameterized for marine mammals and highlight critical research gaps. Large variation exists among available analytical approaches, reflecting the large range of life histories, management needs and data availability across studies. Flexibility in modelling strategy has supported tailored applications to specific case studies but has resulted in limited generality. Despite the many empirical and theoretical uncertainties that remain, bioenergetic models can be used to predict individual and population responses to environmental change and other anthropogenic impacts, thus providing powerful tools to inform effective management and conservation.

This study was supported by Office of Naval Research (ONR) grant N00014- 16-1-2858: “PCoD+: Developing widely-applicable models of the population consequences of disturbance”. Funding support for tagging was provided by the U.S. Navy’s ONR and Living Marine542 Resources (LMR) program, the Chief of Naval Operations’ Energy and Environmental Readiness Division and the NOAA Fisheries Ocean Acoustics Program544 (see Joyce et al. (2020) for details). Hawai‘i field efforts were funded by the U.S. Navy (Pacific Fleet, LMR) and the National Marine Fisheries Service (Pacific Islands Fisheries Science Center). Field efforts were supported by grants from the U.S. Navy’s LMR and N45 programs. Assessing the patterns of wildlife attendance to specific areas is relevant across many fundamental and applied ecological studies, particularly when animals are at risk of being exposed to stressors within or outside the boundaries of those areas. Marine mammals are increasingly being exposed to human activities that may cause behavioral and physiological changes, including military exercises using active sonars. Assessment of the population-level consequences of anthropogenic disturbance requires robust and efficient tools to quantify the levels of aggregate exposure for individuals in a population over biologically relevant time frames. We propose a discrete-space, continuous-time approach to estimate individual transition rates across the boundaries of an area of interest, informed by telemetry data collected with uncertainty. The approach allows inferring the effect of stressors on transition rates, the progressive return to baseline movement patterns, and any difference among individuals. We apply the modeling framework to telemetry data from Blainville's beaked whale (Mesoplodon densirostris) tagged in the Bahamas at the Atlantic Undersea Test and Evaluation Center (AUTEC), an area used by the U.S. Navy for fleet readiness training. We show that transition rates changed as a result of exposure to sonar exercises in the area, reflecting an avoidance response. Our approach supports the assessment of the aggregate exposure of individuals to sonar and the resulting population-level consequences. The approach has potential applications across many applied and fundamental problems where telemetry data are used to characterize animal occurrence within specific areas. Publisher PDF

Funding: K.A.K., R.S.B. and D.P.C. were supported by the E&P Sound and Marine Life Joint Industry Programme (JIP) of the International Association of Oil and Gas Producers (IOGP) (grant no. 00-07-23). K.A.K. was also supported by the National Defense Science and Engineering Graduate (NDSEG) Fellowship. E.P. was supported by the Office of Naval Research (ONR) (grant no. N00014-19-1-2464). D.P.C. was also supported by ONR (grant no. N00014-18-1-2822). Assessing the non-lethal effects of disturbance from human activities is necessary for wildlife conservation and management. However, linking short-term responses to long-term impacts on individuals and populations is a significant hurdle for evaluating the risks of a proposed activity. The Population Consequences of Disturbance (PCoD) framework conceptually describes how disturbance can lead to changes in population dynamics, and its real-world application has led to a suite of quantitative models that can inform risk assessments. Here, we review PCoD models that forecast the possible consequences of a range of disturbance scenarios for marine mammals. In so doing, we identify common themes and highlight general principles to consider when assessing risk. We find that, when considered holistically, these models provide valuable insights into which contextual factors influence a population's degree of exposure and sensitivity to disturbance. We also discuss model assumptions and limitations, identify data gaps and suggest future research directions to enable PCoD models to better inform risk assessments and conservation and management decisions. The general principles explored can help wildlife managers and practitioners identify and prioritize the populations most vulnerable to disturbance and guide industry in planning activities that avoid or mitigate population-level effects. Publisher PDF

Funding: This research was funded by SERDP grant RC20-C2-1097 awarded to Peter Tyack and Prescott Award NA20NMF4390132 awarded to Ashley Barratclough. Epigenetics, specifically DNA methylation, allows for the estimation of animal age from blood or remotely sampled skin. This multi-tissue epigenetic age estimation clock uses 110 longitudinal samples from 34 Navy bottlenose dolphins (Tursiops truncatus), identifying 195 cytosine-phosphate-guanine sites associated with chronological aging via cross-validation with one individual left out in each fold (R2 = 0.95). With a median absolute error of 2.5 years, this clock improves age estimation capacity in wild dolphins, helping conservation efforts and enabling a better understanding of population demographics Publisher PDF

Lightscapes affect life and death of elephant seals during oceanic migrations., Like landscapes of fear, animals are hypothesized to strategically use lightscapes based on intrinsic motivations. However, longitudinal evidence of state-dependent risk aversion has been difficult to obtain in wild animals. Using high-resolution biologgers, we continuously measured body condition, time partitioning, three-dimensional movement, and risk exposure of 71 elephant seals throughout their 7-month foraging migrations (N = 16,000 seal days). As body condition improved from 21 to 32% fat and daylength declined from 16 to 10 hours, seals rested progressively earlier with respect to sunrise, sacrificing valuable nocturnal foraging hours to rest in the safety of darkness. Seals in superior body condition prioritized safety over energy conservation by resting >100 meters deeper where it was 300× darker. Together, these results provide empirical evidence that marine mammals actively use the three-dimensional lightscape to optimize risk-reward trade-offs based on ecological and physiological factors.

Funding: E.P., V.H., J.H., L.N., and D.P.C. were supported by Office of Naval Research (ONR) grant N00014-16-1-2858 (“PCoD+: Developing Widely-Applicable Models of the Population Consequences of Disturbance”). D.P.C. was supported by the Exploration and Production Sound and Marine Life Joint Industry Project of the International Association of Oil and Gas Producers. A.M.d.R. and V.H. were funded by European Research Council Grant no. 322814 awarded to A.M.d.R. M.M. was supported by ONR grant N00014-19-2494. Animals initiate, interrupt, or invest resources in reproduction in light of their physiology and the environment. The energetic risks entailed in an individual’s reproductive strategy can influence the ability to cope with additional stressors, such as anthropogenic climate change and disturbance. To explore the trade-offs between internal state, external resource availability, and reproduction, we applied state-dependent life-history theory (SDLHT) to a dynamic energy budget (DEB) model for long-finned pilot whales (Globicephala melas). We investigated the reproductive strategies emerging from the interplay between fitness maximization and propensity to take energetic risks, as well as the resulting susceptibility of individual vital rates to disturbance. Without disturbance, facultative reproductive behavior from SDLHT and fixed rules in the DEB model led to comparable individual fitness. However, under disturbance, the reproductive strategies emerging from SDLHT increased vulnerability to energetic risks, resulting in lower fitness than fixed rules. These fragile strategies might therefore be unlikely to evolve in the first place. Heterogeneous resource availability favored more cautious (and thus more robust) strategies, particularly when knowledge of resource variation was accurate. Our results demonstrate that the assumptions regarding the dynamic trade-offs underlying an individual’s decision-making can have important consequences for predicting the effects of anthropogenic stressors on wildlife populations. Publisher PDF

Funding: Marine Alliance for Science and Technology for Scotland (MASTs) pooling initiative (LR). MASTs is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions. Ecological niche is traditionally defined at the species level, but individual niches can vary considerably within species. Research on intra-specific niche variation has been focussed on intrinsic drivers. However, differential transmission of socially learned behaviours can also lead to intra-specific niche variation. In sperm whales Physeter macrocephalus, social transmission of information is thought to generate culturally distinct clans, which at times occur sympatrically. Clans have distinct dialects, foraging success rates, and movement patterns, but whether the niches of clan members are also different remains unknown. We evaluated the differences in habitat use of clans off the Galápagos Islands, using data collected over 63 encounters between 1985 and 2014. During encounters, we recorded geographic positions, determined clan identity through analysis of group vocalizations and individual associations, and used topographical and oceanographic variables as proxies of sperm whale prey distribution. We used logistic generalized additive models, fitted with generalized estimating equations to account for spatiotemporal autocorrelation, to predict clan identity as a function of the environment descriptors. Oceanographic variables marginally contributed to differentiating clans. Clan identity could be predicted almost entirely based on geographic location. This fine-scale, within-region spatial partitioning likely derives from whales preferring areas where members of their clans occur over temporal scales of a few months to a few years. By identifying differences in clans’ space use, we have uncovered another level of sperm whale life that is likely influenced by their cultural nature. Postprint

Characterising the spatiotemporal variation of animal behaviour can elucidate the way individuals interact with their environment and allocate energy. Increasing sophistication of tracking technologies paired with novel analytical approaches allows the characterisation of movement dynamics even when an individual is not directly observable. In this study, high-resolution movement data collected via global positioning system (GPS) tracking in three dimensions were paired with topographical information and used in a Bayesian state-space model to describe the flight modes of migrating golden eagles (Aquila chrysaetos) in eastern North America. Our model identified five functional behavioural states, two of which were previously undescribed variations on thermal soaring. The other states comprised gliding, perching and orographic soaring. States were discriminated by movement features in the horizontal (step length and turning angle) and vertical (change in altitude) planes and by the association with ridgelines promoting wind deflection. Tracked eagles spent 2%, 31%, 38%, 9% and 20% of their daytime in directed thermal soaring, gliding, convoluted thermal soaring, perching and orographic soaring, respectively. The analysis of the relative occurrence of these flight modes highlighted yearly, seasonal, age, individual and sex differences in flight strategy and performance. Particularly, less energy-efficient orographic soaring was more frequent in autumn, when thermals were less available. Adult birds were also better at optimising energy efficiency than subadults. Our approach represents the first example of a state-space model for bird flight mode using altitude data in conjunction with horizontal locations and is applicable to other flying organisms where similar data are available. The ability to describe animal movements in a three-dimensional habitat is critical to advance our understanding of the functional processes driving animalsâ decisions. A plain language summary is available for this article.

Between-individual variation in phenotypes within a population is the basis of evolution. However, evolutionary and behavioural ecologists have mainly focused on estimating between-individual variance in mean trait and neglected variation in within-individual variance, or predictability of a trait. In fact, an important assumption of mixed-effects models used to estimate between-individual variance in mean traits is that within-individual residual variance (predictability) is identical across individuals. Individual heterogeneity in the predictability of behaviours is a potentially important effect but rarely estimated and accounted for. We used 11 389 measures of docility behaviour from 1576 yellow-bellied marmots (Marmota flaviventris) to estimate between-individual variation in both mean docility and its predictability. We then implemented a double hierarchical animal model to decompose the variances of both mean trait and predictability into their environmental and genetic components. We found that individuals differed both in their docility and in their predictability of docility with a negative phenotypic covariance. We also found significant genetic variance for both mean docility and its predictability but no genetic covariance between the two. This analysis is one of the first to estimate the genetic basis of both mean trait and within-individual variance in a wild population. Our results indicate that equal within-individual variance should not be assumed. We demonstrate the evolutionary importance of the variation in the predictability of docility and illustrate potential bias in models ignoring variation in predictability. We conclude that the variability in the predictability of a trait should not be ignored, and present a coherent approach for its quantification.

The fish‐eating, type‐C ecotype, killer whale is a top predator in the Ross Sea, Antarctica. Increasing knowledge of this animal's foraging habitats, diet and movement patterns is listed amongst the research priorities adopted under the framework of the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR). To contribute to this goal, satellite transmitters were deployed on 10 type‐C killer whales and skin biopsies were obtained from seven individuals in Terra Nova Bay (Ross Sea) during austral summer (January-February) 2015. Hierarchical switching state-space models (hSSSM) were applied to Argos satellite tracking data to describe the movements of tagged whales, which were then paired with available diving data. Stable isotopes analyses were performed on the biopsy samples to describe the diet. A total of 8,803 Argos locations were available to fit the hSSSM. All whales engaged in potential foraging activity in localized areas along the Ross Sea coastline, followed by uninterrupted travel (i.e. migration) outside Antarctic waters, with no evidence of foraging activity. The pattern of deeper dives matched the occurrence of encamped behaviour indicated by the hSSSM results. The stable isotopes analysis indicated that Antarctic toothfish comprised the largest component (35%) of the prey biomass, raising concerns since this species is targeted by commercial fishery in the Ross Sea Region. These results provide new insights into the ecology of type‐C killer whales in the Ross Sea Region, underlining a potential threat from commercial fishing in the area. Considering the recent establishment of the Ross Sea Region Marine Protected Area, these findings will contribute to the required Research and Monitoring Programme of the Marine Protected Area and provide new empirical evidence to inform conservation measures in the existing Terra Nova Bay Antarctic Special Protected Area

MAC received funding from the Department of Energy and Climate Change (DECC) (now Department for Business, Energy and Industrial Strategy) UK, and the MASTS (Marine Alliance for Science and Technology for Scotland) pooling initiative. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions. Annual surveys were funded by DECC, Scottish Natural Heritage (SNH), Beatrice Offshore Windfarm Ltd., Moray Offshore Renewables Ltd, Marine Scotland, The Crown Estate, Highlands and Islands Enterprise and the Universities of St Andrews and Aberdeen. 1. The efficacy of marine protected areas (MPAs) depends on clear conservation objectives and ecologically meaningful boundaries. The east coast of Scotland bottlenose dolphin population expanded its distributional range during the 1990s beyond the boundaries of the Moray Firth Special Area of Conservation (SAC), originally proposed to contain their core area of distribution. Two decades on, this study assesses the importance for this population of St Andrews Bay and the Tay estuary, 300 km south of the SAC. 2. Photoidentification data from 2009 to 2015 were analysed using mark–recapture models to investigate the proportion of the population that uses St Andrews Bay and the Tay estuary. Habitat models were fitted to bottlenose dolphin presence–absence data to identify areas of high use. 3.The estimated number of dolphins using St Andrews Bay and the Tay estuary during the summer increased from 91 (95% confidence interval 78–106) in 2009 to 114 (95% confidence interval 95–137) in 2015, representing, on average, 52.5% of the total estimated east‐coast population for that period. Spatial mixing of individuals during the summer between St Andrews Bay and the Tay estuary and the Moray Firth SAC was estimated to be a minimum of ~6% per year and ~30% over the study period. The entrance to the Firth of Tay and waters around Montrose were identified as areas of consistent high use. 4. The importance of St Andrews Bay and the Tay estuary reconfirms that effective monitoring of the population requires dedicated effort in both this area and the SAC. The results lead to consideration of the wider context of area‐based management for the conservation/management of highly mobile wide‐ranging species and human activities that might impact them. Postprint

This study was supported by Office of Naval Research grant N00014‐16‐1‐2858: ‘PCoD+: Developing widely‐applicable models of the population consequences of disturbance’. DPC, MM, EAM and LKS were supported by the E&P Sound and Marine Life Joint Industry Project of the International Association of Oil and Gas Producers. JAG was supported by funding from the Young Investigator Program at the Office of Naval Research (award no. N00014‐16‐1‐2477). VH was funded by European Research Council Grant No. 322814 awarded to A.M. de Roos. Animals make behavioural and reproductive decisions that maximise their lifetime reproductive success, and thus their fitness, in light of periodic and stochastic variability of the environment. Modelling the variation of an individual's energy levels formalises this tradeoff and helps to quantify the population‐level consequences of stressors (e.g. disturbance from human activities and environmental change) that can affect behaviour or physiology. In this study, we develop a dynamic state variable model for the spatially explicit behaviour, physiology and reproduction of a female, long‐lived, migratory marine vertebrate. The model can be used to investigate the spatio‐temporal patterns of behaviour and reproduction that allow an individual to maximise its overall reproductive output. We parametrised the model for eastern North Pacific blue whales Balaenoptera musculus, and used it to predict the effects of changing environmental conditions and increasing human disturbance on the population's vital rates. In baseline conditions, the model output had high fidelity to observed energy dynamics, movement patterns and reproductive strategies. Simulated scenarios suggested that environmental changes could have severe consequences on the population's vital rates, but that individuals could tolerate high levels of anthropogenic disturbance. However, this ability depended on where, when and how often disturbance occurred. In scenarios with both environmental change and anthropogenic disturbance, synergistic interactions caused stronger effects than in isolation. In general, larger body size offered a buffer against stochasticity and disturbance, and, consequently, we predicted juveniles to be more susceptible to disturbance. We also predicted that females prioritise their own survival at the expense of the current reproductive attempt, presumably the result of their long lifespan. Our approach provides a general framework to make predictions of the cumulative and synergistic effects of human disturbance and climate change on migratory populations, which can inform effective management and conservation efforts. Publisher PDF

Several species of deep-diving odontocetes, including beaked whales, sperm whales and pilot whales, are associated with steep continental slope habitat in the northeast Atlantic, where they feed on cephalopods and meso-bathypelagic fish. Some species such as sperm whales are listed as vulnerable by the IUCN red list, whilst many beaked whale species are listed as data deficient, therefore, understanding their habitat is key for their successful conservation. A six-day survey was carried out in November 2017 to characterise the habitat of deep-diving odontocete species in this region by sampling multiple aspects of the ecosystem, from oceanographic features through to prey and predator distributions. Sampling of odontocete occurrence was carried out using both visual and acoustic surveys, supported by ten stations that sampled conductivity, temperature, fluorescence and zooplankton; at five of these stations, fish and cephalopod communities were also sampled during dusk and dawn to a maximum depth of 356 m. Results showed that temperature, salinity, and fluorescence were stable throughout the study area. Zooplankton were dominated by copepods, fish were dominated by myctophids, whilst cephalopods were dominated by Todarodes sagittatus and two Histioteuthis spp. Pilot whales, sperm whales and Sowerby's beaked whales were detected both acoustically and visually, along with other delphinids in large numbers, providing evidence that the study covered rich foraging habitat for these species, especially along the 1000 m–2000 m deep continental slope.

The ability to quantify animals’ feeding activity and the resulting changes in their body condition as they move in the environment is fundamental to our understanding of a population’s ecology. We use satellite tracking data from northern elephant seals (Mirounga angustirostris), paired with simultaneous diving information, to develop a Bayesian state-space model that concurrently estimates an individual’s location, feeding activity, and changes in condition. The model identifies important foraging areas and times, the relative amount of feeding occurring therein, and thus the different behavioral strategies in which the seals engage. The fitness implications of these strategies can be assessed by looking at the resulting variation in individuals’ condition, which in turn affects the condition and survival of their offspring. Therefore, our results shed light on the processes affecting an individual’s decision-making as it moves and feeds in the environment. In addition, we demonstrate how the model can be used to simulate realistic patterns of disturbance at different stages of the trip, and how the predicted accumulation of lipid reserves varies as a consequence. Particularly, disturbing an animal in periods of high feeding activity or shortly after leaving the colony was predicted to have the potential to lead to starvation. In contrast, an individual could compensate even for very severe disturbance if such disturbance occurred outside the main foraging grounds. Our modeling approach is applicable to marine mammal species that perform drift dives and can be extended to other species where an individual’s buoyancy can be inferred from its diving behavior.

Managing the nonlethal effects of disturbance on wildlife populations has been a long‐term goal for decision makers, managers, and ecologists, and assessment of these effects is currently required by European Union and United States legislation. However, robust assessment of these effects is challenging. The management of human activities that have nonlethal effects on wildlife is a specific example of a fundamental ecological problem: how to understand the population‐level consequences of changes in the behavior or physiology of individual animals that are caused by external stressors. In this study, we review recent applications of a conceptual framework for assessing and predicting these consequences for marine mammal populations. We explore the range of models that can be used to formalize the approach and we identify critical research gaps. We also provide a decision tree that can be used to select the most appropriate model structure given the available data. Synthesis and applications: The implementation of this framework has moved the focus of discussion of the management of nonlethal disturbances on marine mammal populations away from a rhetorical debate about defining negligible impact and toward a quantitative understanding of long‐term population‐level effects. Here we demonstrate the framework's general applicability to other marine and terrestrial systems and show how it can support integrated modeling of the proximate and ultimate mechanisms that regulate trait‐mediated, indirect interactions in ecological communities, that is, the nonconsumptive effects of a predator or stressor on a species' behavior, physiology, or life history.

Whale watching can affect cetacean behaviour, and can in some cases lead to long-term negative effects on survival and reproduction. The waters of Juneau (Alaska) represent a summer feeding ground for the Central North Pacific stock of humpback whales, Megaptera novaeangliae (Borowski, 1781). The recent dramatic expansion of the local whale-watching industry has raised concerns over the potential negative impact of such activity on the whales. The aim of this study was to understand the impact of whale-watching vessels on humpback whale behaviour in Juneau. From June to September 2015, land-based observations were conducted to record whale behaviour in the presence and absence of vessels. Markov chains and model simulations showed that the presence of vessels was not associated with a reduction in the time spent feeding. Whales exhibited more surface-active behaviour and travelled less in the presence of vessels. The lack of an overt response of feeding whales to whale-watching disturbance could result from the importance of the Southeast Alaska study region as a feeding ground, which could make the whales less apt to interrupt feeding under disturbance; however, The high proportion of travelling observed may indicate that the study area represents a travelling corridor between other, localized feeding areas. The increase in surface-active behaviour may represent a response to vessel presence that could potentially lead to energetic consequences. The large number of whale-watching vessels operating could lead to the increased tolerance of whales towards whale-watching activities. The lack of responses in this study may suggest that the reactions of whales are context specific, depending on their previous experience with vessels and their foraging behaviour, for example. Subtler negative effects could still occur, however, and a prolonged monitoring programme is required to assess less overt behavioural responses of whales over time.

Integrating behavior and physiology is critical to formulating new hypotheses on the evolution of animal life-history strategies. Migratory capital breeders acquire most of the energy they need to sustain migration, gestation, and lactation before parturition. Therefore, when predicting the impact of environmental variation on such species, a mechanistic understanding of the physiology of their migratory behavior is required. Using baleen whales as a model system, we developed a dynamic state variable model that captures the interplay among behavioral decisions, energy, reproductive needs, and the environment. We applied the framework to blue whales (Balaenoptera musculus) in the eastern North Pacific Ocean and explored the effects of environmental and anthropogenic perturbations on female reproductive success. We demonstrate the emergence of migration to track prey resources, enabling us to quantify the trade-offs among capital breeding, body condition, and metabolic expenses. We predict that periodic climatic oscillations affect reproductive success less than unprecedented environmental changes do. The effect of localized, acute anthropogenic impacts depended on whales' behavioral response to the disturbance; chronic, but weaker, disturbances had little effect on reproductive success. Because we link behavior and vital rates by modeling individuals' energetic budgets, we provide a general framework to investigate the ecology of migration and assess the population consequences of disturbance, while identifying critical knowledge gaps.

Assessments of anthropogenic impacts on marine wildlife often concentrate on large-scale displacement. However, changes in the activity patterns of animals that do not flee could also affect their energy balance. Increasing boat traffic raises concerns for exposed marine mammals. Understanding risk perception is critical for effective mitigation and management, but it is hard to disentangle the effect of noise, physical presence of boats, and context. We used passive acoustic techniques to quantify how boat disturbance affected bottlenose dolphin foraging activity, and characterized the conditions influencing responses. To account for potential masking effects of boat noise on foraging vocalizations (buzzes), we developed a novel procedure to estimate the relationship between buzz detection probability and noise levels in particular 1/3 octave bands. Bayesian hierarchical modeling was then used to assess the effect of boat presence on buzz occurrence, as well as potential interactions with noise level, location, year, day, hour, dolphin group size, and boat type and number. Our results indicate that boat presence was associated with a short-term 49% reduction in foraging activity, but there was no relationship with noise level. Differences between sites and between years suggested a variable susceptibility depending on foraging conditions. This effect increased for increasing number of boats and depended on boat type. This is the first study conclusively showing that boat physical presence, and not just noise, plays a large role in disturbance.

Human activities can influence the movement of organisms, either repelling or attracting individuals depending on whether they interfere with natural behavioural patterns or enhance access to food. To discern the processes affecting such interactions, an appropriate analytical approach must reflect the motivations driving behavioural decisions at multiple scales. In this study, we developed a modelling framework for the analysis of foraging trips by central place foragers. By recognising the distinction between movement phases at a larger scale and movement steps at a finer scale, our model can identify periods when animals are actively following moving attractors in their landscape. We applied the framework to GPS tracking data of northern fulmars Fulmarus glacialis, paired with contemporaneous fishing boat locations, to quantify the putative scavenging activity of these seabirds on discarded fish and offal. We estimated the rate and scale of interaction between individual birds and fishing boats and the interplay with other aspects of a foraging trip. The model classified periods when birds were heading out to sea, returning towards the colony or following the closest boat. The probability of switching towards a boat declined with distance and varied depending on the phase of the trip. The maximum distance at which a bird switched towards the closest boat was estimated around 35 km, suggesting the use of olfactory information to locate food. Individuals spent a quarter of a foraging trip, on average, following fishing boats, with marked heterogeneity among trips and individuals. Our approach can be used to characterise interactions between central place foragers and different anthropogenic or natural stimuli. The model identifies the processes influencing central place foraging at multiple scales, which can improve our understanding of the mechanisms underlying movement behaviour and characterise individual variation in interactions with a range of human activities that may attract or repel these species. Therefore, it can be adapted to explore the movement of other species that are subject to multiple dynamic drivers.

Mediterranean fin whales comprise a genetically distinct population, listed as Vulnerable (VU) in the IUCN Red List. Collisions with vessels are believed to represent the main cause of human-induced mortality. The identification of critical habitats (including migration routes) incorporating satellite telemetry data is therefore crucial to develop focussed conservation efforts. Between 2012 and 2015 thirteen fin whales were equipped with satellite transmitters, 8 in the Pelagos Sanctuary (although two ceased within two days) and 5 in the Strait of Sicily, to evaluate movements and habitat use. A hierarchical switching state-space model was used to identify transiting and area-restricted search (ARS) behaviours, believed to indicate foraging activities. All whales undertook mid- to long-distance migrations, crossing some of the world’s busiest maritime routes. Areas where the animals predominantly engaged in ARS behaviour were identified in both study areas. The telemetry data were compared with results from ecosystem niche modelling, and showed that 80% of tagged whale positions was near (

This project was funded through the DECC Offshore Energy Strategic Environmental Assessment Programme using equipment previously purchased by DECC, Scottish Government, Oil and Gas UK, COWRIE and Moray Offshore Renewables Ltd. Scottish Natural Heritage, Beatrice Offshore Windfarm Ltd., MORL, Marine Scotland, The Crown Estate and Highlands and Islands Enterprise all provided funding for photo-identification surveys. Data have been made available in the Dryad repository at https://doi.org/10.5061/dryad.g3603. The development of risk assessments for the exposure of protected populations to noise from coastal construction is constrained by uncertainty over the nature and extent of marine mammal responses to man-made noise. Stakeholder concern often focuses on the potential for local displacement caused by impact piling, where piles are hammered into the seabed. To mitigate this threat, use of vibration piling, where piles are shaken into place with a vibratory hammer, is often encouraged due to presumed impact reduction. However, data on comparative responses of cetaceans to these different noise sources are lacking. We studied the responses of bottlenose dolphins and harbor porpoises to both impact and vibration pile driving noise during harbor construction works in northeast Scotland, using passive acoustic monitoring devices to record cetacean activity and noise recorders to measure and predict received noise levels. Local abundance and patterns of occurrence of bottlenose dolphins were also compared with a five-year baseline. The median peak-to-peak source level estimated for impact piling was 240 dB re 1 μPa (single-pulse sound exposure level [SEL] 198 dB re 1 μPa2 s), and the r.m.s. source level for vibration piling was 192 dB re 1 μPa. Predicted received broadband SEL values 812 m from the piling site were markedly lower due to high propagation loss: 133.4 dB re 1 μPa2 s (impact) and 128.9 dB re 1 μPa2 s (vibration). Bottlenose dolphins and harbor porpoises were not excluded from sites in the vicinity of impact piling or vibration piling; nevertheless, some small effects were detected. Bottlenose dolphins spent a reduced period of time in the vicinity of construction works during both impact and vibration piling. The probability of occurrence of both cetacean species was also slightly less during periods of vibration piling. This work provides developers and managers with the first evidence of the comparative effects of vibration and impact piling on small cetaceans, enabling more informed risk assessments, policy frameworks, and mitigation plans. In particular, our results emphasize the need for better understanding of noise levels and behavioral responses to vibration piling before recommending its use to mitigate impact piling. Publisher PDF

Summary 1. Individuals make behavioural decisions by weighing potential advantages and costs (e.g. increased food intake vs. increased risk of predation). When animals change their activities in response to a perceived threat, their energetic input may decline. Marine ecotourism, including whale and dolphin watching, is growing globally and cetaceans perceive interactions with tour vessels as a form of risk. Observable behavioural changes need to be linked to bioenergetic effects to determine the potential population consequences of this disturbance. 2. We developed a theoretical optimal dive model for bottlenose dolphins under three potential types of perceived risk resulting from human interactions at the surface (decreasing instantaneous risk, increasing instantaneous risk and no risk). We compared the predictions of these theoretical models to observed dive cycles of foraging male and female dolphins in the presence and absence of tour vessels. We used mixture models to classify dive types and mixed effects models to analyse changes in the interbreath interval of surface and bottom dives and the frequency of estimated bottom dives. 3. Males significantly increased bottom time and performed fewer bottom dives when boats were present, matching predictions of our theoretical model for perceived decreasing instantaneous risk. In contrast, females significantly decreased bottom times and increased the frequency of bottom dives, matching predictions from the model for perceived increasing instantaneous risk. Therefore, our empirical results suggest differences in the perception of risk between sexes. 4. Synthesis and applications. By comparing theoretical predictions with observed dive data, our study suggests that boat interactions during foraging can cause decreased net energy gain over a foraging bout for both sexes, with females being more impacted. The population under study is currently listed as critically endangered. Understanding whether these predicted energetic impacts affect an individual’s vital rates will provide a link to the population-level consequences of this disturbance. Previous analytical approaches have failed to capture the costs associated with disturbance during foraging, leading to management recommendations that only protect animals from increased energetic expenditure. We suggest that the current management scheme should be revised to include foraging areas in order to secure the energy intake of animals.

Appropriate management of the effects of human activities on animal populations requires quantification of the rate at which animals encounter stressors. Such activities are heterogeneously distributed in space, as are the individual animals in a population. This will result in a heterogeneous exposure rate, which is also likely to vary over time. A spatially explicit analysis of individual exposure is therefore required. We applied Bayesian spatially explicit capture‐recapture models to photo-identification data to estimate the home range of well-marked individuals in a protected coastal population of bottlenose dolphins. Model results were combined with the estimated distribution of boat traffic to quantify how exposure to this disturbance varied in time and space. Variability in exposure between individuals was also investigated using a mixed-effects model. The cumulative individual exposure to boat traffic varied between summers, depending both on the overall area usage and the degree of individual movement around the activity centres. Despite this variability, regions of higher risk could be identified. There were marked inter-individual differences in the predicted amount of time dolphins spent in the presence of boats, and individuals tended to be consistently over- or underexposed across summers. Our study offers a framework to describe the temporal, spatial and individual variation in exposure to anthropogenic stressors when individuals can be repeatedly identified over time. It provides opportunities to map exposure risk and understand how this evolves in time at both individual and population levels. The outcome of such modelling can be used as a robust evidence base to support management decisions.

The potential impacts of underwater noise on marine mammals are widely recognised, but uncertainty over variability in baseline noise levels often constrains efforts to manage these impacts. This paper characterises natural and anthropogenic contributors to underwater noise at two sites in the Moray Firth Special Area of Conservation, an important marine mammal habitat that may be exposed to increased shipping activity from proposed offshore energy developments. We aimed to establish a pre-development baseline, and to develop ship noise monitoring methods using Automatic Identification System (AIS) and time-lapse video to record trends in noise levels and shipping activity. Our results detail the noise levels currently experienced by a locally protected bottlenose dolphin population, explore the relationship between broadband sound exposure levels and the indicators proposed in response to the EU Marine Strategy Framework Directive, and provide a ship noise assessment toolkit which can be applied in other coastal marine environments.

1. Common bottlenose dolphins Tursiops truncatus interact with fish farms in the Mediterranean Sea. These interactions were investigated in a Greek bay by incorporating multiple geographic, bathymetric, oceanographic, and anthropogenic variables. 2. Generalized additive models (GAMs) and generalized estimation equations (GEEs) were used to describe dolphin presence. Visual surveys were conducted over 2909 km under favourable viewing conditions that included 54 dolphin group follows for 457 km. Sea surface temperature (SST) and chlorophyll-a (Chl-a) data were obtained from remote sensing imagery, and distances to sources of human influences including fish farms, a ferro-nickel plant, and a slag disposal area were calculated within a geographic information system (GIS). 3. Bottlenose dolphins were encountered mainly in the south-eastern portion of the study area, and occurrence was not clearly related to SST and Chl-a, nor the ferro-nickel plant or nearby slag disposal area. 4. Dolphin occurrence generally increased within 20 km of fish farms, with four farms and dolphins displaying a positive relationship, seven no clear relationship, and two a negative one. 5. While it is likely that uneaten food and other detritus attract dolphin prey, individual farms (or clusters of farms) clearly had a different appeal. The proximity of the ferro-nickel plant and slag disposal area to ‘attractive’ fish farms could compromise dolphin health, but physiological data are unavailable. 6. The modelling of multiple variables allowed for a description of dolphin habitat use and attraction to some fish farms. More such data analysed in similar manner would be instructive for other areas where marine mammals and fish farms co-occur. Copyright © 2013 John Wiley & Sons, Ltd.

Summary 1. Understanding which environmental factors drive foraging preferences is critical for the development of effective management measures, but resource use patterns may emerge from processes that occur at different spatial and temporal scales. Direct observations of foraging are also especially challenging in marine predators, but passive acoustic techniques provide opportunities to study the behaviour of echolocating species over a range of scales. 2. We used an extensive passive acoustic data set to investigate the distribution and temporal dynamics of foraging in bottlenose dolphins using the Moray Firth (Scotland, UK). Echolocation buzzes were identified with a mixture model of detected echolocation inter-click intervals and used as a proxy of foraging activity. A robust modelling approach accounting for autocorrelation in the data was then used to evaluate which environmental factors were associated with the observed dynamics at two different spatial and temporal scales. 3. At a broad scale, foraging varied seasonally and was also affected by seabed slope and shelf-sea fronts. At a finer scale, we identified variation in seasonal use and local interactions with tidal processes. Foraging was best predicted at a daily scale, accounting for site specificity in the shape of the estimated relationships. 4. This study demonstrates how passive acoustic data can be used to understand foraging ecology in echolocating species and provides a robust analytical procedure for describing spatiotemporal patterns. Associations between foraging and environmental characteristics varied according to spatial and temporal scale, highlighting the need for a multi-scale approach. Our results indicate that dolphins respond to coarser scale temporal dynamics, but have a detailed understanding of finer-scale spatial distribution of resources.

This paper presents the empirical probability density of the power spectral density as a tool to assess the field performance of passive acoustic monitoring systems and the statistical distribution of underwater noise levels across the frequency spectrum. Using example datasets, it is shown that this method can reveal limitations such as persistent tonal components and insufficient dynamic range, which may be undetected by conventional techniques. The method is then combined with spectral averages and percentiles, which illustrates how the underlying noise level distributions influence these metrics. This combined approach is proposed as a standard, integrative presentation of ambient noise spectra.

Sperm whale Physeter macrocephalus habitat preferences are still poorly understood inthe Mediterranean, despite the population being classified as ‘Endangered’ by the IUCN. Techniquesto make the best use of multiple data sources are important in improving this situation. This work pro-vides a detailed evaluation of sperm whale distribution and habitat use around the Balearic Islandsusing a novel analytical framework that combines transect and follow data while accounting for anyautocorrelation present. During dedicated research cruises (2003 to 2008), sperm whales werelocated by listening at regular intervals along a search track and subsequently followed acoustically.Sperm whales were encountered 56 times and followed for periods ranging from a few hours to 3 d.Logistic Generalized Additive Models were used to model the probability of whale presence acrossthe study area as a function of environmental variables, and Generalized Estimating Equations wereused to account for autocorrelation. The results suggest that sperm whales do not use the region uni-formly and that topography plays a key role in shaping their distribution. Moreover, solitary individ-uals were found to use the habitat differently from groups. This segregation appeared to be driven bywater temperature and might reflect different needs or intraspecific competition. By shedding lighton sperm whale habitat preference in such a critical area, this study represents an important steptowards the implementation of conservation measures for this population.KEY WORDS: Habitat modelling · Cetacean · Distribution · Balearic Islands · GAM · GEE · Multi-scale

We review recent work that developed new techniques for underwater noise assessment that integrate acoustic monitoring with automatic identification system (AIS) shipping data and time-lapse video, meteorological, and tidal data. Two sites were studied within the Moray Firth Special Area of Conservation (SAC) for bottlenose dolphins, where increased shipping traffic is expected from construction of offshore wind farms outside the SAC. Noise exposure varied markedly between the sites, and natural and anthropogenic contributions were characterized using multiple data sources. At one site, AIS-operating vessels accounted for total cumulative sound exposure (0.1–10 kHz), suggesting that noise modeling using the AIS would be feasible.

We review recent work that developed new techniques for underwater noise assessment that integrate acoustic monitoring with automatic identification system (AIS) shipping data and time-lapse video, meteorological, and tidal data. Two sites were studied within the Moray Firth Special Area of Conservation (SAC) for bottlenose dolphins, where increased shipping traffic is expected from construction of offshore wind farms outside the SAC. Noise exposure varied markedly between the sites, and natural and anthropogenic contributions were characterized using multiple data sources. At one site, AIS-operating vessels accounted for total cumulative sound exposure (0.1-10 kHz), suggesting that noise modeling using the AIS would be feasible.

This work received funding from the Marine Alliance for Science and Technology for Scotland (MASTS pooling initiative). Human activities that impact wildlife do not necessarily remove individuals from populations. They may also change individual behaviour in ways that have sublethal effects. This has driven interest in developing analytical tools that predict the population consequences of short-term behavioural responses. In this study, we incorporate empirical information on the ecology of a population of bottlenose dolphins into an individual-based model that predicts how individuals' behavioural dynamics arise from their underlying motivational states, as well as their interaction with boat traffic and dredging activities. We simulate the potential effects of proposed coastal developments on this population and predict that the operational phase may affect animals' motivational states. For such results to be relevant for management, the effects on individuals' vital rates also need to be quantified. We investigate whether the relationship between an individual's exposure and the survival of its calves can be directly estimated using a Bayesian multi-stage model for calf survival. The results suggest that any effect on calf survival is probably small and that a significant relationship could only be detected in large, closely studied populations. Our work can be used to guide management decisions, accelerate the consenting process for coastal and offshore developments and design targeted monitoring Postprint