Deborah Jill Fraser Russell

Combining individual animal movement and ancillary biotelemetry data to investigate population-level activity budgets

Recent technological advances have permitted the collection of detailed animal location and ancillary biotelemetry data that facilitate inference about animal movement and associated behaviors. However, these rich sources of individual information, location, and biotelemetry data, are typically analyzed independently, with population-level inferences remaining largely post hoc. We describe a hierarchical modeling approach, which is able to integrate location and ancillary biotelemetry (e.g., physiological or accelerometer) data from many individuals. We can thus obtain robust estimates of (1) population-level movement parameters and (2) activity budgets for a set of behaviors among which animals transition as they respond to changes in their internal and external environment. Measurement error and missing data are easily accommodated using a state-space formulation of the proposed hierarchical model. Using Bayesian analysis methods, we demonstrate our modeling approach with location and dive activity data from 17 harbor seals (Phoca vitulina) in the United Kingdom. Based jointly on movement and diving activity, we identified three distinct movement behavior states: resting, foraging, and transit, and estimated population-level activity budgets to these three states. Because harbor seals are known to dive for both foraging and transit (but not usually for resting), we compared these results to a similar population- level analysis utilizing only location data. We found that a large proportion of time steps were mischaracterized when behavior states were inferred from horizontal trajectory alone, with 33% of time steps exhibiting a majority of dive activity assigned to the resting state. Only 1% of these time steps were assigned to resting when inferred from both trajectory and dive activity data using our integrated modeling approach. There is mounting evidence of the potential perils of inferring animal behavior based on trajectory alone, but there fortunately now exist many flexible analytical techniques for extracting more out of the increasing wealth of information afforded by recent advances in biologging technology.

Marine mammals trace anthropogenic structures at sea

On land, species from all trophic levels have adapted to fill vacant niches in environments heavily modified by humans (e.g. [1]). In the marine environment, ocean infrastructure has led to artificial reefs, resulting in localized increases in fish and crustacean density [2]. Whether marine apex predators exhibit behavioural adaptations to utilise such a scattered potential resource is unknown. Using high resolution GPS data we show how infrastructure, including wind turbines and pipelines, shapes the movements of individuals from two seal species (Phoca vitulina and Halichoerus grypus). Using state-space models, we infer that these animals are using structures to forage. We highlight the ecological consequences of such behaviour, at a time of unprecedented developments in marine infrastructure.

Assessing performance of Bayesian state-space models fit to Argos Satellite telemetry locations processed with Kalman Filtering

Argos recently implemented a new algorithm to calculate locations of satellite-tracked animals that uses a Kalman filter (KF). The KF algorithm is reported to increase the number and accuracy of estimated positions over the traditional Least Squares (LS) algorithm, with potential advantages to the application of state-space methods to model animal movement data. We tested the performance of two Bayesian state-space models (SSMs) fitted to satellite tracking data processed with KF algorithm. Tracks from 7 harbour seals (Phoca vitulina) tagged with ARGOS satellite transmitters equipped with Fastloc GPS loggers were used to calculate the error of locations estimated from SSMs fitted to KF and LS data, by comparing those to “true” GPS locations. Data on 6 fin whales (Balaenoptera physalus) were used to investigate consistency in movement parameters, location and behavioural states estimated by switching state-space models (SSSM) fitted to data derived from KF and LS methods. The model fit to KF locations improved the accuracy of seal trips by 27% over the LS model. 82% of locations predicted from the KF model and 73% of locations from the LS model were <5 km from the corresponding interpolated GPS position. Uncertainty in KF model estimates (5.6±5.6 km) was nearly half that of LS estimates (11.6±8.4 km). Accuracy of KF and LS modelled locations was sensitive to precision but not to observation frequency or temporal resolution of raw Argos data. On average, 88% of whale locations estimated by KF models fell within the 95% probability ellipse of paired locations from LS models. Precision of KF locations for whales was generally higher. Whales’ behavioural mode inferred by KF models matched the classification from LS models in 94% of the cases. State-space models fit to KF data can improve spatial accuracy of location estimates over LS models and produce equally reliable behavioural estimates.

Carer provisioning rules in an obligate cooperative breeder: prey type, size and delivery rate

Providing food to developing offspring is beneficial for offspring but costly for carers. Understanding patterns of provisioning thus yields important insights into how selection shapes (allo-) parental care strategies. Broadly, offspring development will be influenced by three components of provisioning (prey type, size and delivery rate). However, all three variables are rarely considered simultaneously, leading to suggestions that the results of many studies are misleading. Additionally, few studies have examined the provisioning strategies of breeders and non-breeding helpers in obligate cooperative breeders, wherein reproduction without help is typically unsuccessful. We investigated these components of provisioning in obligately cooperative chestnut-crowned babblers (Pomatostomus ruficeps). Prey type was associated with size, and delivery rate was the best predictor of the overall amount of food provided by carers. As broods aged, breeders and helpers similarly modified the relative proportion of different prey provided and increased both prey size and delivery rate. Breeding females contributed less prey than male breeders and adult helpers, and were the only carers to load-lighten by reducing their provisioning rates in the presence of additional carers. While our results suggest that breeders and helpers follow broadly comparable provisioning rules, they are also consistent with the idea that, in obligately cooperative species, breeding females benefit more from conserving resources for future reproduction than do helpers which have a low probability of breeding independently.

Uncovering the links between foraging and breeding regions in a highly mobile mammal

1. The annual cycle of many animals is characterized by the need to satisfy different life history priorities, often requiring seasonal movements. For such species, investigating carryover effects (such as the year-long drivers of breeding success) and managing protected areas effectively, relies on quantifying these movements. Here, we model the seasonal movements of the UK population of grey seals Halichoerus grypus and show how insights from the model can improve its management. 2. We fit a hidden process model to two types of information – regional population redistribution and individual movements – to estimate the seasonal transition probabilities of breeding female grey seals among four regions around the UK. 3. We found that between 21% and 58% of females used different regions for breeding and foraging. 4. For our study period, we detected an increase in the breeding performance of animals that foraged in the Hebrides and South-East Coast. 5. Grey seal Special Areas of Conservation (SACs) were designed to encompass a significant proportion of the UK breeding population: ~ 40% of the breeding females in our study area. Of the females breeding on SACs, only 15% breed in Northern Scotland, but up to 50% forage there. Our results indicate that, by only considering the breeding distribution of females that breed in SACs, the impact of anthropogenic activities on nearby SACs may be overestimated, whereas impacts on remote SACs may be underestimated. 6. Synthesis and applications. By quantifying the link between the foraging and breeding distributions of grey seals, management of breeding populations can be focused on the foraging regions where the resources necessary for reproduction are acquired. The construction of marine developments is dependent on demonstrating that they will not have an adverse effect on the integrity of Special Areas of Conservation (SACs), and we have shown that this requires consideration of the seasonal transition probabilities estimated in this study. Our specific results provide support for management strategies that jointly consider SACs and Marine Protected Areas (MPAs). More generally, we prescribe combinations of data on population size, breeding performance and individual movement that can enable our framework to be applied to seasonally migrating species.

Avoidance of wind farms by harbour seals is limited to pile driving activities

Summary As part of global efforts to reduce dependence on carbon‐based energy sources there has been a rapid increase in the installation of renewable energy devices. The installation and operation of these devices can result in conflicts with wildlife. In the marine environment, mammals may avoid wind farms that are under construction or operating. Such avoidance may lead to more time spent travelling or displacement from key habitats. A paucity of data on at‐sea movements of marine mammals around wind farms limits our understanding of the nature of their potential impacts. Here, we present the results of a telemetry study on harbour seals Phoca vitulina in The Wash, south‐east England, an area where wind farms are being constructed using impact pile driving. We investigated whether seals avoid wind farms during operation, construction in its entirety, or during piling activity. The study was carried out using historical telemetry data collected prior to any wind farm development and telemetry data collected in 2012 during the construction of one wind farm and the operation of another. Within an operational wind farm, there was a close‐to‐significant increase in seal usage compared to prior to wind farm development. However, the wind farm was at the edge of a large area of increased usage, so the presence of the wind farm was unlikely to be the cause. There was no significant displacement during construction as a whole. However, during piling, seal usage (abundance) was significantly reduced up to 25 km from the piling activity; within 25 km of the centre of the wind farm, there was a 19 to 83% (95% confidence intervals) decrease in usage compared to during breaks in piling, equating to a mean estimated displacement of 440 individuals. This amounts to significant displacement starting from predicted received levels of between 166 and 178 dB re 1 μPa(p‐p). Displacement was limited to piling activity; within 2 h of cessation of pile driving, seals were distributed as per the non‐piling scenario. Synthesis and applications. Our spatial and temporal quantification of avoidance of wind farms by harbour seals is critical to reduce uncertainty and increase robustness in environmental impact assessments of future developments. Specifically, the results will allow policymakers to produce industry guidance on the likelihood of displacement of seals in response to pile driving; the relationship between sound levels and avoidance rates; and the duration of any avoidance, thus allowing far more accurate environmental assessments to be carried out during the consenting process. Further, our results can be used to inform mitigation strategies in terms of both the sound levels likely to cause displacement and what temporal patterns of piling would minimize the magnitude of the energetic impacts of displacement.

Sound exposure in harbour seals during the installation of an offshore wind farm: predictions of auditory damage

Summary 1. With ambitious renewable energy targets, pile driving associated with offshore wind farm construction will become widespread in the marine environment. Many proposed wind farms overlap with the distribution of seals, and sound from pile driving has the potential to cause auditory damage. 2. We report on a behavioural study during the construction of a wind farm using data from GPS/GSM tags on 24 harbour seals Phoca vitulina L. Pile driving data and acoustic propagation models, together with seal movement and dive data, allowed the prediction of auditory damage in each seal. 3. Growth and recovery functions for auditory damage were combined to predict temporary auditory threshold shifts in each seal. Further, M-weighted cumulative sound exposure levels [cSELs(Mpw)] were calculated and compared to permanent auditory threshold shift exposure criteria for pinnipeds in water exposed to pulsed sounds. 4. The closest distance of each seal to pile driving varied from 4� 7t o 40 � 5 km, and predicted maximum cSELs(Mpw) ranged from 170� 7 to 195� 3d B re 1lPa 2 -s for individual seals. Comparison to exposure criteria suggests that half of the seals exceeded estimated permanent auditory damage thresholds. 5. Prediction of auditory damage in marine mammals is a rapidly evolving field and has a number of key uncertainties associated with it. These include how sound propagates in shallow water environments and the effects of pulsed sounds on seal hearing; as such, our predictions should be viewed in this context. 6. Policy implications. We predicted that half of the tagged seals received sound levels from pile driving that exceeded auditory damage thresholds for pinnipeds. These results have implications for offshore industry and will be important for policymakers developing guidance for pile driving. Developing engineering solutions to reduce sound levels at source or methods to deter animals from damage risk zones, or changing temporal patterns of piling could potentially reduce auditory damage risk. Future work should focus on validating these predictions by collecting auditory threshold information pre- and post-exposure to pile driving. Ultimately, information on population-level impacts of exposure to pile driving is required to ensure that offshore industry is developed in an environmentally sustainable manner.

Navigating uncertain waters: a critical review of inferring foraging behaviour from location and dive data in pinnipeds

In the last thirty years, the emergence and progression of biologging technology has led to great advances in marine predator ecology. Large databases of location and dive observations from biologging devices have been compiled for an increasing number of diving predator species (such as pinnipeds, sea turtles, seabirds and cetaceans), enabling complex questions about animal activity budgets and habitat use to be addressed. Central to answering these questions is our ability to correctly identify and quantify the frequency of essential behaviours, such as foraging. Despite technological advances that have increased the quality and resolution of location and dive data, accurately interpreting behaviour from such data remains a challenge, and analytical methods are only beginning to unlock the full potential of existing datasets. This review evaluates both traditional and emerging methods and presents a starting platform of options for future studies of marine predator foraging ecology, particularly from location and two-dimensional (time-depth) dive data. We outline the different devices and data types available, discuss the limitations and advantages of commonly-used analytical techniques, and highlight key areas for future research. We focus our review on pinnipeds - one of the most studied taxa of marine predators - but offer insights that will be applicable to other air-breathing marine predator tracking studies. We highlight that traditionally-used methods for inferring foraging from location and dive data, such as first-passage time and dive shape analysis, have important caveats and limitations depending on the nature of the data and the research question. We suggest that more holistic statistical techniques, such as state-space models, which can synthesise multiple track, dive and environmental metrics whilst simultaneously accounting for measurement error, offer more robust alternatives. Finally, we identify a need for more research to elucidate the role of physical oceanography, device effects, study animal selection, and developmental stages in predator behaviour and data interpretation.

Addressing uncertainty in marine resource management; combining community engagement and tracking technology to characterise human behavior

Small-scale fisheries provide an essential source of food and employment for coastal communities, yet the availability of detailed information on the spatiotemporal distribution of fishing effort to support resource management at a country level is scarce. Here, using a national-scale study in the Republic of Congo, we engaged with fishers from 23 of 28 small-scale fisheries landing sites along the coast to demonstrate how combining community engagement and relatively low cost Global Positioning System (GPS) trackers can rapidly provide fine-scale information on: (1) the behavioural dynamics of the fishers and fleets that operate within this sector; and (2) the location, size and attributes of important fishing grounds upon which communities are dependent. This multi-disciplinary approach should be considered within a global context where uncertainty over the behaviour of marine and terrestrial resource-users can lead to management decisions that potentially compromise local livelihoods, conservation, and resource sustainability goals. This article is protected by copyright. All rights reserved

Dynamic habitat corridors for marine predators; intensive use of a coastal channel by harbour seals is modulated by tidal currents

Previous studies have found that predators utilise habitat corridors to ambush prey moving through them. In the marine environment, coastal channels effectively act as habitat corridors for prey movements, and sightings of predators in such areas suggest that they may target these for foraging. Unlike terrestrial systems where the underlying habitat structure is generally static, corridors in marine systems are in episodic flux due to water movements created by tidal processes. Although these hydrographic features can be highly complex, there is generally a predictable underlying cyclic tidal pattern to their structure. For marine predators that must find prey that is often patchy and widely distributed, the underlying temporal predictability in potential foraging opportunities in marine corridors may be important drivers in their use. Here, we used data from land-based sightings and 19 harbour seals (Phoca vitulina) tagged with high-resolution GPS telemetry to investigate the spatial and temporal distribution patterns of seals in a narrow tidal channel. These seals showed a striking pattern in their distribution; all seals spent a high proportion of their time around the narrowest point of the channel. There was also a distinctive tidal pattern in the use of the channel; sightings of seals in the water peaked during the flood tide and were at a minimum during the ebb tide. This pattern is likely to be related to prey availability and/or foraging efficiency driven by the underlying tidal pattern in the water movements through the channel.Significance StatementTo maximise foraging efficiency, predators often make use of narrow constrictions in habitat to intercept prey using these corridors for movement. In the marine environment, narrow channels may act as corridors, and sightings of predators suggest that they may target these for foraging. Despite this, there is little information on how individual predators use such areas. Here, we investigate how individual harbour seals use a narrow coastal channel subject to strong tidal currents; results showed that seals spent the majority of their time at the narrowest point of the channel foraging during peak tidal currents. This highlights the importance of narrow channels for marine predators and suggests that this usually wide-ranging predator may restrict its geographic range to forage in the channel as a result of increased prey availability and/or foraging efficiency driven by water movements through the narrow corridor.