Variation in habitat preference and distribution of harbour porpoises west of Scotland

The waters off the west coast of Scotland have one of the highest densities of harbour porpoise (Phocoena phocoena) in Europe. Harbour porpoise are listed under Annex II of the EU Habitats Directive, requiring the designation of Special Areas of Conservation (SACs) for the species’ protection and conservation. The main aim of this thesis is to identify habitat preferences for harbour porpoise, and key regions that embody these preferences, which could therefore be suitable as SACs; and to determine how harbour porpoise use these regions over time and space. Designed visual and acoustic line-transect surveys were conducted between 2003 and 2008. Generalised Estimating Equations (GEEs) were used to determine relationships between the relative density of harbour porpoise and temporally and spatially variable oceanographic covariates. Predictive models showed that depth, slope, distance to land and spring tidal range were all important in explaining porpoise distribution. There were also significant temporal variations in habitat use. However, whilst some variation was observed among years and months, consistent preferences for water depths between 50 and 150 m and highly sloped regions were observed across the temporal models. Predicted surfaces revealed a consistent inshore distribution for the species throughout the west coast of Scotland. Regional models revealed similar habitat preferences to the full-extent models, and indicated that the Small Isles and Sound of Jura were the most consistently important regions for harbour porpoise, and that these regions could be suitable as SACs. The impacts of seal scarers on distribution and habitat use were also investigated, and there were indications that these devices have the potential to displace harbour porpoise. These results should be considered in the assessment of sites for SAC designation, and in implementing appropriate conservation measures for harbour porpoise

Methods for monitoring for the population consequences of disturbance in marine mammals : a review

This work was sponsored by the Office of Naval Research: Marine Mammal Biology Program, under award N000141612858.Assessing the non-lethal effects of disturbance and their population-level consequences is a significant ecological and conservation challenge, because it requires extensive baseline knowledge of behavioral patterns, life-history and demography. However, for many marine mammal populations, this knowledge is currently lacking and it may take decades to fill the gaps. During this time, undetected population declines may occur. In this study we identify methods that can be used to monitor populations subject to disturbance and provide insights into the processes through which disturbance may affect them. To identify and address the knowledge gaps highlighted above, we reviewed the literature to identify suitable response variables and methods for monitoring these variables. We also used existing models of the population consequences of disturbance (PCoD) to identify demographic characteristics (e.g., the proportion of immature animals in the population, or the ratio of calves/pups to mature females) that may be strongly correlated with population status and therefore provide early warnings of future changes in abundance. These demographic characteristics can be monitored using established methods such as visual surveys combined with photogrammetry, and capture-recapture analysis. Individual health and physiological variables can also inform PCoD assessment and can be monitored using photogrammetry, remote tissue sampling, hands-on assessment and individual tracking. We then conducted a workshop to establish the relative utility and feasibility of all these approaches for different groups of marine mammal species. We describe how future marine mammal monitoring programs can be designed to inform population-level analysis.Publisher PDFPeer reviewe

Combining bioenergetics and movement models to improve understanding of the population consequences of disturbance

Funding: Naval Facilities Engineering Systems Command EXWC - N3943019C2175.We developed dynamic bioenergetics models to investigate how behavioural responses to anthropogenic disturbance events might affect the population dynamics of three marine mammal species (harbour porpoise, grey seal and harbour seal) with contrasting life-history traits (capital versus income breeders) and movement behaviour (resident versus nomadic). We used these models to analyse how individual vital rates were affected by differences in the probability of disturbance and the duration of any behavioural response, while taking account of uncertainty in the model parameters and heterogeneity in behaviour. The outputs of individual movement models and telemetry data were then used to determine how the probability of exposure might vary among species, individuals, and geographical locations. We then demonstrate how these estimated probabilities of exposure can be translated into probabilities of disturbance. For illustrative purposes, we modelled the potential effects of a temporary decrease in energy assimilation associated with a series of disturbance events that might realistically occur during the construction of an offshore windfarm. Offspring starvation mortality was the vital rate that was most affected by these disturbance events. Monitoring of rate should be considered as standard practice so that populations responses can be detected as early as possible. Predicted effects on individual vital rates depended on the species' movement behaviour and the likely density of animals where the modelled construction activity was assumed to take place. The magnitude of these effects also depended critically on the assumed duration of the reduction in energy assimilation. No direct estimates of this variable are currently available, but we suggest some ways in which it could be estimated. The described approach could be extended to other species and activities, given sufficient information to parameterise the component models. However, we emphasise the need to account for among-individual heterogeneities and uncertainties in the values of the many model parameters.Publisher PDFPeer reviewe

Population consequences of the Deepwater Horizon oil spill on pelagic cetaceans

This research was made possible by a grant from the Gulf of Mexico Research Initiative to the Consortium for Advanced Research on Marine Mammal Health Assessment (CARMMHA). T.A.M. acknowledges partial support by CEAUL (funded by FCT−Fundação para a Ciência e a Tecnologia, Portugal, through project UIDB/00006/2020).The Deepwater Horizon disaster resulted in the release of 490000 m3 of oil into the northern Gulf of Mexico. We quantified population consequences for pelagic cetaceans, including sperm whales, beaked whales and 11 species of delphinids. We used existing spatial density models to establish pre-spill population size and distribution, and overlaid an oil footprint to estimate the proportion exposed to oil. This proportion ranged from 0.058 (Atlantic spotted dolphin, 95% CI = 0.041-0.078) to 0.377 (spinner dolphin, 95% CI = 0.217-0.555). We adapted a population dynamics model, developed for an estuarine population of bottlenose dolphins, to each pelagic species by scaling demographic parameters using literature-derived estimates of gestation duration. We used expert elicitation to translate knowledge from dedicated studies of oil effects on bottlenose dolphins to pelagic species and address how density dependence may affect reproduction. We quantified impact by comparing population trajectories under baseline and oil-impacted scenarios. The number of lost cetacean years (difference between trajectories, summed over years) ranged from 964 (short-finned pilot whale, 95% CI = 385-2291) to 32584 (oceanic bottlenose dolphin, 95% = CI 13377-71967). Maximum proportional population decrease ranged from 1.3% (Atlantic spotted dolphin 95% CI = 0.5-2.3) to 8.4% (spinner dolphin 95% CI = 3.2-17.7). Estimated time to recover to 95% of baseline was >10 yr for spinner dolphin (12 yr, 95% CI = 0-21) and sperm whale (11 yr, 95% CI = 0-21), while 7 taxonomic units remained within 95% of the baseline population size (time to recover, therefore, as per its definition, was 0). We investigated the sensitivity of results to alternative plausible inputs. Our methods are widely applicable for estimating population effects of stressors in the absence of direct measurements.Publisher PDFPeer reviewe

Quantifying the age structure of free-ranging delphinid populations : testing the accuracy of Unoccupied Aerial System photogrammetry

This study was funded by NOAA-PIFSC and RCUH JIMAR (NA19NMF4720181, NA16NMF4320058), CIMAR (NA21NMF4320043), and the Office of Naval Research (N000142012624).Understanding the population health status of long-lived and slow-reproducing species is critical for their management. However, it can take decades with traditional monitoring techniques to detect population-level changes in demographic parameters. Early detection of the effects of environmental and anthropogenic stressors on vital rates would aid in forecasting changes in population dynamics and therefore inform management efforts. Changes in vital rates strongly correlate with deviations in population growth, highlighting the need for novel approaches that can provide early warning signs of population decline (e.g., changes in age structure). We tested a novel and frequentist approach, using Unoccupied Aerial System (UAS) photogrammetry, to assess the population age structure of small delphinids. First, we measured the precision and accuracy of UAS photogrammetry in estimating total body length (TL) of trained bottlenose dolphins (Tursiops truncatus). Using a log-transformed linear model, we estimated TL using the blowhole to dorsal fin distance (BHDF) for surfacing animals. To test the performance of UAS photogrammetry to age-classify individuals, we then used length measurements from a 35-year dataset from a free-ranging bottlenose dolphin community to simulate UAS estimates of BHDF and TL. We tested five age classifiers and determined where young individuals (Publisher PDFPeer reviewe

Resolving issues with environmental impact assessment of marine renewable energy installations

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 PDFPeer reviewe

Variation in habitat preference and distribution of harbour porpoises west of Scotland

The waters off the west coast of Scotland have one of the highest densities of harbour porpoise (Phocoena phocoena) in Europe. Harbour porpoise are listed under Annex II of the EU Habitats Directive, requiring the designation of Special Areas of Conservation (SACs) for the species’ protection and conservation. The main aim of this thesis is to identify habitat preferences for harbour porpoise, and key regions that embody these preferences, which could therefore be suitable as SACs; and to determine how harbour porpoise use these regions over time and space. Designed visual and acoustic line-transect surveys were conducted between 2003 and 2008. Generalised Estimating Equations (GEEs) were used to determine relationships between the relative density of harbour porpoise and temporally and spatially variable oceanographic covariates. Predictive models showed that depth, slope, distance to land and spring tidal range were all important in explaining porpoise distribution. There were also significant temporal variations in habitat use. However, whilst some variation was observed among years and months, consistent preferences for water depths between 50 and 150 m and highly sloped regions were observed across the temporal models. Predicted surfaces revealed a consistent inshore distribution for the species throughout the west coast of Scotland. Regional models revealed similar habitat preferences to the full-extent models, and indicated that the Small Isles and Sound of Jura were the most consistently important regions for harbour porpoise, and that these regions could be suitable as SACs. The impacts of seal scarers on distribution and habitat use were also investigated, and there were indications that these devices have the potential to displace harbour porpoise. These results should be considered in the assessment of sites for SAC designation, and in implementing appropriate conservation measures for harbour porpoise.EThOS - Electronic Theses Online ServiceGBUnited Kingdo

A sampling, exposure and receptor framework for identifying factors that modulate behavioural responses to disturbance in cetaceans

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 PDFPeer reviewe

Context-dependent variability in the predicted daily energetic costs of disturbance for blue whales

Funding: This study was supported by the Office of Naval Research (ONR) [grant number N00014–19-1-2464: “BRS4PCoD:Integrating the results of Behavioral Response Studies intomodels of the Population Consequences of Disturbance”]. J.A.G., D.E.C. and J.A.F. were supported by the National Science Foundation (Division of Integrative Organismal Systems) [grant number 1656691], ONR Young Investigator Program [grant number N000141612477], ONR Defense University Research Instrumentation Program [grant number N000141612546] and Stanford University’s Terman and Bass Fellowships. The SOCAL-BRS project was supported by the US Navy’s Chief of Naval Operations Environmental Readiness Division, the US Navy’s Living Marine Resources Program and the Marine Mammal Program of the Office of Naval Research.Assessing the long-term consequences of sub-lethal anthropogenic disturbance on wildlife populations requires integrating data on fine-scale individual behavior and physiology into spatially and temporally broader, population-level inference. A typical behavioral response to disturbance is the cessation of foraging, which can be translated into a common metric of energetic cost. However, this necessitates detailed empirical information on baseline movements, activity budgets, feeding rates and energy intake, as well as the probability of an individual responding to the disturbance-inducing stressor within different exposure contexts. Here, we integrated data from blue whales (Balaenoptera musculus) experimentally exposed to military active sonar signals with fine-scale measurements of baseline behavior over multiple days or weeks obtained from accelerometry loggers, telemetry tracking and prey sampling. Specifically, we developed daily simulations of movement, feeding behavior and exposure to localized sonar events of increasing duration and intensity and predicted the effects of this disturbance source on the daily energy intake of an individual. Activity budgets and movements were highly variable in space and time and among individuals, resulting in large variability in predicted energetic intake and costs. In half of our simulations, an individual’s energy intake was unaffected by the simulated source. However, some individuals lost their entire daily energy intake under brief or weak exposure scenarios. Given this large variation, population-level models will have to assess the consequences of the entire distribution of energetic costs, rather than only consider single summary statistics. The shape of the exposure-response functions also strongly influenced predictions, reinforcing the need for contextually explicit experiments and improved mechanistic understanding of the processes driving behavioral and physiological responses to disturbance. This study presents a robust approach for integrating different types of empirical information to assess the effects of disturbance at spatio-temporal and ecological scales that are relevant to management and conservation.Publisher PDFPeer reviewe