Kylie L. Scales

REVIEW: On the Front Line: frontal zones as priority at‐sea conservation areas for mobile marine vertebrates

Summary 1. Identifying priority areas for marine vertebrate conservation is complex because species of conservation concern are highly mobile, inhabit dynamic habitats and are difficult to monitor. 2. Many marine vertebrates are known to associate with oceanographic fronts – physical interfaces at the transition between water masses – for foraging and migration, making them important candidate sites for conservation. Here, we review associations between marine vertebrates and fronts and how they vary with scale, regional oceanography and foraging ecology. 3. Accessibility, spatiotemporal predictability and relative productivity of front-associated foraging habitats are key aspects of their ecological importance. Predictable mesoscale (10s– 100s km) regions of persistent frontal activity (‘frontal zones’) are particularly significant. 4. Frontal zones are hotspots of overlap between critical habitat and spatially explicit anthropogenic threats, such as the concentration of fisheries activity. As such, they represent tractable conservation units, in which to target measures for threat mitigation. 5. Front mapping via Earth observation (EO) remote sensing facilitates identification and monitoring of these hotspots of vulnerability. Seasonal or climatological products can locate biophysical hotspots, while near-real-time front mapping augments the suite of tools supporting spatially dynamic ocean management. 6. Synthesis and applications. Frontal zones are ecologically important for mobile marine vertebrates. We surmise that relative accessibility, predictability and productivity are key biophysical characteristics of ecologically significant frontal zones in contrasting oceanographic regions. Persistent frontal zones are potential priority conservation areas for multiple marine vertebrate taxa and are easily identifiable through front mapping via EO remote sensing. These insights are useful for marine spatial planning and marine biodiversity conservation, both within Exclusive Economic Zones and in the open oceans.

A Bird’s Eye View of Discard Reforms: Bird-Borne Cameras Reveal Seabird/Fishery Interactions

Commercial capture fisheries produce huge quantities of offal, as well as undersized and unwanted catch in the form of discards. Declines in global catches and legislation to ban discarding will significantly reduce discards, but this subsidy supports a large scavenger community. Understanding the potential impact of declining discards for scavengers should feature in an eco-system based approach to fisheries management, but requires greater knowledge of scavenger/fishery interactions. Here we use bird-borne cameras, in tandem with GPS loggers, to provide a unique view of seabird/fishery interactions. 20,643 digital images (one min−1) from ten bird-borne cameras deployed on central place northern gannets Morus bassanus revealed that all birds photographed fishing vessels. These were large (>15 m) boats, with no small-scale vessels. Virtually all vessels were trawlers, and gannets were almost always accompanied by other scavenging birds. All individuals exhibited an Area-Restricted Search (ARS) during foraging, but only 42% of ARS were associated with fishing vessels, indicating much ‘natural’ foraging. The proportion of ARS behaviours associated with fishing boats were higher for males (81%) than females (30%), although the reasons for this are currently unclear. Our study illustrates that fisheries form a very important component of the prey-landscape for foraging gannets and that a discard ban, such as that proposed under reforms of the EU Common Fisheries Policy, may have a significant impact on gannet behaviour, particularly males. However, a continued reliance on ‘natural’ foraging suggests the ability to switch away from scavenging, but only if there is sufficient food to meet their needs in the absence of a discard subsidy.

Mesoscale fronts as foraging habitats: composite front mapping reveals oceanographic drivers of habitat use for a pelagic seabird

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

Ocean-wide tracking of pelagic sharks reveals extent of overlap with longline fishing hotspots

Significance Shark populations are declining worldwide because of overexploitation by fisheries with unknown consequences for ecosystems. Although the harvest of oceanic sharks remains largely unregulated, knowing precisely where they interact with fishing vessels will better aid their conservation. We satellite track six species of shark and two entire longline fishing vessel fleets across the North Atlantic over multiple years. Sharks actively select and aggregate in space-use “hotspots” characterized by thermal fronts and high productivity. However, longline fishing vessels also target these habitats and efficiently track shark movements seasonally, leading to an 80% spatial overlap. Areas of highest overlap between sharks and fishing vessels show persistence between years, suggesting current hotspots are at risk, and arguing for introduction of international catch limits. Overfishing is arguably the greatest ecological threat facing the oceans, yet catches of many highly migratory fishes including oceanic sharks remain largely unregulated with poor monitoring and data reporting. Oceanic shark conservation is hampered by basic knowledge gaps about where sharks aggregate across population ranges and precisely where they overlap with fishers. Using satellite tracking data from six shark species across the North Atlantic, we show that pelagic sharks occupy predictable habitat hotspots of high space use. Movement modeling showed sharks preferred habitats characterized by strong sea surface-temperature gradients (fronts) over other available habitats. However, simultaneous Global Positioning System (GPS) tracking of the entire Spanish and Portuguese longline-vessel fishing fleets show an 80% overlap of fished areas with hotspots, potentially increasing shark susceptibility to fishing exploitation. Regions of high overlap between oceanic tagged sharks and longliners included the North Atlantic Current/Labrador Current convergence zone and the Mid-Atlantic Ridge southwest of the Azores. In these main regions, and subareas within them, shark/vessel co-occurrence was spatially and temporally persistent between years, highlighting how broadly the fishing exploitation efficiently “tracks” oceanic sharks within their space-use hotspots year-round. Given this intense focus of longliners on shark hotspots, our study argues the need for international catch limits for pelagic sharks and identifies a future role of combining fine-scale fish and vessel telemetry to inform the ocean-scale management of fisheries.

Scale of inference: on the sensitivity of habitat models for wide‐ranging marine predators to the resolution of environmental data

&NA; Understanding and predicting the responses of wide‐ranging marine predators such as cetaceans, seabirds, sharks, turtles, pinnipeds and large migratory fish to dynamic oceanographic conditions requires habitat‐based models that can sufficiently capture their environmental preferences. Marine ecosystems are inherently dynamic, and animal–environment interactions are known to occur over multiple, nested spatial and temporal scales. The spatial resolution and temporal averaging of environmental data layers are therefore key considerations in modelling the environmental determinants of habitat selection. The utility of environmental data contemporaneous to animal presence or movement (e.g. daily, weekly), versus synoptic products (monthly, seasonal, climatological) is currently debated, as are the trade‐offs between near real‐time, high resolution and composite (i.e. synoptic, cloud‐free) data fields. Using movement simulations with built‐in environmental preferences in combination with both modelled and remotely‐sensed (ROMS, MODIS‐Aqua) sea surface temperature (SST) fields, we explore the effects of spatial and temporal resolution (3–111 km, daily–climatological) in predictive habitat models. Results indicate that models fitted using seasonal or climatological data fields can introduce bias in presence‐availability designs based upon animal movement datasets, particularly in highly dynamic oceanographic domains. These effects were pronounced where models were constructed using seasonal or climatological fields of coarse (> 0.25 degree) spatial resolution. However, cloud obstruction can lead to significant information loss in remotely‐sensed data fields. We found that model accuracy decreased substantially above 70% data loss. In cloudy regions, weekly or monthly environmental data fields may therefore be preferable. These findings have important implications for marine resource management, particularly in identifying key habitats for populations of conservation concern, and in forecasting climate‐mediated ecosystem changes.

Basking sharks and oceanographic fronts: quantifying associations in the north‐east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.

GPS tracking reveals rafting behaviour of Northern Gannets (Morus bassanus): implications for foraging ecology and conservation

Capsule Three quarters of tracked Northern Gannets (Morus bassanus) at Grassholm gathered in rafts around the colony, concentrated within a recently designated at-sea Special Protection Area (SPA), but rafting was not correlated with foraging effort. Aims To investigate the incidence, distribution and foraging implications of Northern Gannet rafting behaviour in waters adjacent to a large colony. Methods Using bird-borne global positioning system (GPS) loggers we reconstructed at-sea movement and used a speed filter to identify rafting behaviour within 10 km of the colony. We mapped the spatial distribution of rafting events from 160 breeding individuals over 5 years, and investigated the relationship between foraging effort (trip duration and total distance travelled) and the presence/absence of rafting. Results On average, 74% of tracked birds engaged in rafting. Of the 381 foraging trips analysed, rafting was recorded on 237 (62%). Birds were more likely to raft on outbound (224 trips, 59%), than inbound journeys (38 trips, 10%). Presence/absence of rafting did not correlate significantly with foraging trip distance or duration nor with duration of nest attendance. The majority of rafting was concentrated in a 2-km radius around the colony within a recently designated seaward SPA extension. Birds showed low individual repeatability in rafting, although there was lower variation within, than among, individuals. Conclusion Our results show that rafting is important for breeding gannets on Grassholm, and a recently designated at-sea SPA encapsulates the core distribution of rafting. Rafting did not appear to be correlated with foraging behaviour. Given the dearth of literature on rafting and the wealth of GPS tracking data for seabirds, we suggest that similar research be conducted elsewhere to further elucidate the ecological and applied significance of this behaviour.

A dynamic ocean management tool to reduce bycatch and support sustainable fisheries

Dynamic management approaches protect endangered bycatch species but with much greater efficiency than existing static closures. Seafood is an essential source of protein for more than 3 billion people worldwide, yet bycatch of threatened species in capture fisheries remains a major impediment to fisheries sustainability. Management measures designed to reduce bycatch often result in significant economic losses and even fisheries closures. Static spatial management approaches can also be rendered ineffective by environmental variability and climate change, as productive habitats shift and introduce new interactions between human activities and protected species. We introduce a new multispecies and dynamic approach that uses daily satellite data to track ocean features and aligns scales of management, species movement, and fisheries. To accomplish this, we create species distribution models for one target species and three bycatch-sensitive species using both satellite telemetry and fisheries observer data. We then integrate species-specific probabilities of occurrence into a single predictive surface, weighing the contribution of each species by management concern. We find that dynamic closures could be 2 to 10 times smaller than existing static closures while still providing adequate protection of endangered nontarget species. Our results highlight the opportunity to implement near real-time management strategies that would both support economically viable fisheries and meet mandated conservation objectives in the face of changing ocean conditions. With recent advances in eco-informatics, dynamic management provides a new climate-ready approach to support sustainable fisheries.

Integrating Dynamic Subsurface Habitat Metrics Into Species Distribution Models

Species distribution models (SDMs) have become key tools for describing and predicting species habitats. In the marine domain, environmental data used in modelling species distributions are often remotely sensed, and as such have limited capacity for interpreting the vertical structure of the water column, or are sampled in situ, offering minimal spatial and temporal coverage. Advances in ocean models have improved our capacity to explore subsurface ocean features, yet there has been limited integration of such features in SDMs. Using output from a data-assimilative configuration of the Regional Ocean Modeling System, we examine the effect of including dynamic subsurface variables in SDMs to describe the habitats of four pelagic predators in the California Current System (swordfish Xiphias gladius, blue sharks Prionace glauca, common thresher sharks Alopias vulpinus, and shortfin mako sharks Isurus oxyrinchus). Species data were obtained from the California Drift Gillnet observer program (1997-2017). We used boosted regression trees to explore the incremental improvement enabled by dynamic subsurface variables that quantify the structure and stability of the water column: isothermal layer depth and bulk buoyancy frequency. The inclusion of these dynamic subsurface variables significantly improved model explanatory power for most species. Model predictive performance also significantly improved, but only for species that had strong affiliations with dynamic variables (swordfish and shortfin mako sharks) rather than static variables (blue sharks and common thresher sharks). Geospatial predictions for all species showed the integration of isothermal layer depth and bulk buoyancy frequency contributed value at the mesoscale level (<100 km) and varied spatially throughout the study domain. These results highlight the utility of including dynamic subsurface variables in SDM development and support the continuing ecological use of biophysical output from ocean circulation models.

Multi-year tracking reveals extensive pelagic phase of juvenile loggerhead sea turtles in the North Pacific

BackgroundThe juvenile stage of loggerhead sea turtles (Caretta caretta) can last for decades. In the North Pacific Ocean, much is known about their seasonal movements in relation to pelagic habitat, yet understanding their multi-year, basin-scale movements has proven more difficult. Here, we categorize the large-scale movements of 231 turtles satellite tracked from 1997 to 2013 and explore the influence of biological and environmental drivers on basin-scale movement.ResultsResults show high residency of juvenile loggerheads within the Central North Pacific and a moderate influence of the Earth’s magnetic field, but no real-time environmental driver to explain migratory behavior.ConclusionsWe suggest the Central North Pacific acts as important developmental foraging grounds for young juvenile loggerhead sea turtles, rather than just a migratory corridor. We propose several hypotheses that may influence the connectivity between western and eastern juvenile loggerhead foraging grounds in the North Pacific Ocean.