Identifying predictable foraging habitats for a wide-ranging marine predator using ensemble ecological niche models

Aim: Ecological niche modelling can provide valuable insight into species' environmental preferences and aid the identification of key habitats for populations of conservation concern. Here, we integrate biologging, satellite remote-sensing and ensemble ecological niche models (EENMs) to identify predictable foraging habitats for a globally important population of the grey-headed albatross (GHA) Thalassarche chrysostoma. Location: Bird Island, South Georgia; Southern Atlantic Ocean. Methods: GPS and geolocation-immersion loggers were used to track at-sea movements and activity patterns of GHA over two breeding seasons (n = 55; brood-guard). Immersion frequency (landings per 10-min interval) was used to define foraging events. EENM combining Generalized Additive Models (GAM), MaxEnt, Random Forest (RF) and Boosted Regression Trees (BRT) identified the biophysical conditions characterizing the locations of foraging events, using time-matched oceanographic predictors (Sea Surface Temperature, SST; chlorophyll a, chl-a; thermal front frequency, TFreq; depth). Model performance was assessed through iterative cross-validation and extrapolative performance through cross-validation among years. Results: Predictable foraging habitats identified by EENM spanned neritic (<500 m), shelf break and oceanic waters, coinciding with a set of persistent biophysical conditions characterized by particular thermal ranges (3–8 °C, 12–13 °C), elevated primary productivity (chl-a > 0.5 mg m−3) and frequent manifestation of mesoscale thermal fronts. Our results confirm previous indications that GHA exploit enhanced foraging opportunities associated with frontal systems and objectively identify the APFZ as a region of high foraging habitat suitability. Moreover, at the spatial and temporal scales investigated here, the performance of multi-model ensembles was superior to that of single-algorithm models, and cross-validation among years indicated reasonable extrapolative performance. Main conclusions: EENM techniques are useful for integrating the predictions of several single-algorithm models, reducing potential bias and increasing confidence in predictions. Our analysis highlights the value of EENM for use with movement data in identifying at-sea habitats of wide-ranging marine predators, with clear implications for conservation and management.

Identifying predictable foraging habitats for a wide-ranging marine predator using ensemble ecological niche models

Aim: Ecological niche modelling can provide valuable insight into species' environmental preferences and aid the identification of key habitats for populations of conservation concern. Here, we integrate biologging, satellite remote-sensing and ensemble ecological niche models (EENMs) to identify predictable foraging habitats for a globally important population of the grey-headed albatross (GHA) Thalassarche chrysostoma. Location: Bird Island, South Georgia; Southern Atlantic Ocean. Methods: GPS and geolocation-immersion loggers were used to track at-sea movements and activity patterns of GHA over two breeding seasons (n = 55; brood-guard). Immersion frequency (landings per 10-min interval) was used to define foraging events. EENM combining Generalized Additive Models (GAM), MaxEnt, Random Forest (RF) and Boosted Regression Trees (BRT) identified the biophysical conditions characterizing the locations of foraging events, using time-matched oceanographic predictors (Sea Surface Temperature, SST; chlorophyll a, chl-a; thermal front frequency, TFreq; depth). Model performance was assessed through iterative cross-validation and extrapolative performance through cross-validation among years. Results: Predictable foraging habitats identified by EENM spanned neritic (<500 m), shelf break and oceanic waters, coinciding with a set of persistent biophysical conditions characterized by particular thermal ranges (3–8 °C, 12–13 °C), elevated primary productivity (chl-a > 0.5 mg m−3) and frequent manifestation of mesoscale thermal fronts. Our results confirm previous indications that GHA exploit enhanced foraging opportunities associated with frontal systems and objectively identify the APFZ as a region of high foraging habitat suitability. Moreover, at the spatial and temporal scales investigated here, the performance of multi-model ensembles was superior to that of single-algorithm models, and cross-validation among years indicated reasonable extrapolative performance. Main conclusions: EENM techniques are useful for integrating the predictions of several single-algorithm models, reducing potential bias and increasing confidence in predictions. Our analysis highlights the value of EENM for use with movement data in identifying at-sea habitats of wide-ranging marine predators, with clear implications for conservation and management.

Plastic and marine turtles: a review and call for research

Plastic debris is now ubiquitous in the marine environment affecting a wide range of taxa, from microscopic zooplankton to large vertebrates. Its persistence and dispersal throughout marine ecosystems has meant that sensitivity toward the scale of threat is growing, particularly for species of conservation concern, such as marine turtles. Their use of a variety of habitats, migratory behaviour, and complex life histories leave them subject to a host of anthropogenic stressors, including exposure to marine plastic pollution. Here, we review the evidence for the effects of plastic debris on turtles and their habitats, highlight knowledge gaps, and make recommendations for future research. We found that, of the seven species, all are known to ingest or become entangled in marine debris. Ingestion can cause intestinal blockage and internal injury, dietary dilution, malnutrition, and increased buoyancy which in turn can result in poor health, reduced growth rates and reproductive output, or death. Entanglement in plastic debris (including ghost fishing gear) is known to cause lacerations, increased drag—which reduces the ability to forage effectively or escape threats—and may lead to drowning or death by starvation. In addition, plastic pollution may impact key turtle habitats. In particular, its presence on nesting beaches may alter nest properties by affecting temperature and sediment permeability. This could influence hatchling sex ratios and reproductive success, resulting in population level implications. Additionally, beach litter may entangle nesting females or emerging hatchlings. Lastly, as an omnipresent and widespread pollutant, plastic debris may cause wider ecosystem effects which result in loss of productivity and implications for trophic interactions. By compiling and presenting this evidence, we demonstrate that urgent action is required to better understand this issue and its effects on marine turtles, so that appropriate and effective mitigation policies can be developed.

A review of climate change and the implementation of marine biodiversity legislation in the United Kingdom

1. Marine legislation, the key means by which the conservation of marine biodiversity is achieved, has been developing since the 1960s. In recent decades, an increasing focus on ‘holistic’ policy development is evident, compared with earlier ‘piecemeal’ sectoral approaches. Important marine legislative tools being used in the United Kingdom, and internationally, include the designation of marine protected areas and the Marine Strategy Framework Directive (MSFD) with its aim of meeting ‘Good Environmental Status’ (GES) for European seas by 2020. 2. There is growing evidence of climate change impacts on marine biodiversity, which may compromise the effectiveness of any legislation intended to promote sustainable marine resource management. 3. A review of key marine biodiversity legislation relevant to the UK shows climate change was not considered in the drafting of much early legislation. Despite the huge increase in knowledge of climate change impacts in recent decades, legislation is still limited in how it takes these impacts into account. There is scope, however, to account for climate change in implementing much of the legislation through (a) existing references to environmental variability; (b) review cycles; and (c) secondary legislation and complementary policy development. 4. For legislation relating to marine protected areas (e.g. the EC Habitats and Birds Directives), climate change has generally not been considered in the site-designation process, or for ongoing management, with the exception of the Marine (Scotland) Act. Given that changing environmental conditions (e.g. rising temperatures and ocean acidification) directly affect the habitats and species that sites are designated for, how this legislation is used to protect marine biodiversity in a changing climate requires further consideration. 5. Accounting for climate change impacts on marine biodiversity in the development and implementation of legislation is vital to enable timely, adaptive management responses. Marine modelling can play an important role in informing management decisions.

Analyses of sublittoral macrobenthic community change in a marine nature reserve using similarity profiles (SIMPROF).

Sublittoral macrobenthic communities in the Skomer Marine Nature Reserve (SMNR), Pembrokeshire, Wales, were sampled at 10 stations in 1993, 1996, 1998, 2003, 2007 and 2009 using a Day grab and a 0.5 mm mesh. The time series is analysed using Similarities Profiles (SIMPROF) tests and associated methods. Q-mode analysis using clustering with Type 1 SIMPROF addresses multivariate structure among samples, showing that there is clear structure associated with differences among years. Inverse (r-mode) analysis using Type 2 SIMPROF decisively rejects a hypothesis that species are not associated with each other. Clustering of the variables (species) with Type 3 SIMPROF identifies groups of species which covary coherently through the time-series. The time-series is characterised by a dramatic decline in abundances and diversity between the 1993 and 1996 surveys. By 1998 there had been a shift in community composition from the 1993 situation, with different species dominating. Communities had recovered in terms of abundance and species richness, but different species dominated the community. No single factor could be identified which unequivocally explained the dramatic changes observed in the SMNR. Possible causes were the effects of dispersed oil and dispersants from the Sea Empress oil spill in February 1996 and the cessation of dredge-spoil disposal off St Anne’s Head in 1995, but the most likely cause was severe weather. With many species, and a demonstrable recovery from an impact, communities within the SMNR appear to be diverse and resilient. If attributable to natural storms, the changes observed here indicate that natural variability may be much more important than is generally taken into account in the design of monitoring programmes.

Relationships between biodiversity and the stability of marine ecosystems: comparisons at a European scale using meta-analysis.

The relationship between biodiversity and stability of marine benthic assemblages was investigated using existing data sets (n = 28) covering various spatial (m-km) and temporal (1973-2006) scales in different benthic habitats (emergent rock, rock pools and sedimentary habitats) through meta-analyses. Assemblage stability was estimated by measuring temporal variances of species richness, total abundance (density or % cover) and community species composition and abundance structure (using multivariate analyses). Positive relationships between temporal variability in species number and richness were generally observed at both quadrat (<1 m2) and site (100 m2) scales, while no relationships were observed by multivariate analyses. Positive relationships were also observed at the scale of site between temporal variability in species number and variability in community structure with evenness estimates. This implies that the relationship between species richness or evenness and species richness variability is slightly positive and depends on the scale of observation, suggesting that biodiversity per se is important for the stability of ecosystems. Changes within community assemblages in terms of structure are, however, generally independent of biodiversity, suggesting no effect of diversity, but the potential impact of individual species, and/or environmental factors. Except for sedimentary and rock pool habitats, no relationship was observed between temporal variation of the aggregated variable of total abundances and diversity at either scale. Overall our results emphasise that relationships depend on scale of measurements, type of habitats and the marine systems (North Atlantic and Mediterranean) considered.

Assessing the sensitivity of seabed species and biotopes - the Marine Life Information Network (MarLIN).

Science-based approaches to support the conservation of marine biodiversity have been developed in recent years. They include measures of ‘rarity’, ‘diversity’, ‘importance’, biological indicators of water ‘quality’ and measures of ‘sensitivity’. Identifying the sensitivity of species and biotopes, the main topic of this contribution, relies on accessing and interpreting available scientific data in a structured way and then making use of information technology to disseminate suitably presented information to decision makers. The Marine Life Information Network (MarLIN) has achieved that research for a range of environmentally critical species and biotopes over the past four years and has published the reviews on the MarLIN Web site (www.marlin.ac.uk). Now, by linking the sensitivity database and databases of survey information, sensitivity mapping approaches using GIS are being developed. The methods used to assess sensitivity are described and the approach is advocated for wider application in Europe.

Projections of change in key ecosystem indicators for planning and management of marine protected areas: An example study for European seas

Marine Protected Areas (MPAs) are widely used as tools to maintain biodiversity, protect habitats and ensure that development is sustainable. If MPAs are to maintain their role into the future it is important for managers to understand how conditions at these sites may change as a result of climate change and other drivers, and this understanding needs to extend beyond temperature to a range of key ecosystem indicators. This case study demonstrates how spatially-aggregated model results for multiple variables can provide useful projections for MPA planners and managers. Conditions in European MPAs have been projected for the 2040s using unmitigated and globally managed scenarios of climate change and river management, and hence high and low emissions of greenhouse gases and riverborne nutrients. The results highlight the vulnerability of potential refuge sites in the north-west Mediterranean and the need for careful monitoring at MPAs to the north and west of the British Isles, which may be affected by changes in Atlantic circulation patterns. The projections also support the need for more MPAs in the eastern Mediterranean and Adriatic Sea, and can inform the selection of sites.

Projections of change in key ecosystem indicators for planning and management of marine protected areas: An example study for European seas

Marine Protected Areas (MPAs) are widely used as tools to maintain biodiversity, protect habitats and ensure that development is sustainable. If MPAs are to maintain their role into the future it is important for managers to understand how conditions at these sites may change as a result of climate change and other drivers, and this understanding needs to extend beyond temperature to a range of key ecosystem indicators. This case study demonstrates how spatially-aggregated model results for multiple variables can provide useful projections for MPA planners and managers. Conditions in European MPAs have been projected for the 2040s using unmitigated and globally managed scenarios of climate change and river management, and hence high and low emissions of greenhouse gases and riverborne nutrients. The results highlight the vulnerability of potential refuge sites in the north-west Mediterranean and the need for careful monitoring at MPAs to the north and west of the British Isles, which may be affected by changes in Atlantic circulation patterns. The projections also support the need for more MPAs in the eastern Mediterranean and Adriatic Sea, and can inform the selection of sites.

Application of computer-aided tomography techniques to visualize kelp holdfast structure reveals the importance of habitat complexity for supporting marine biodiversity

Ecosystem engineers that increase habitat complexity are keystone species in marine systems, increasing shelter and niche availability, and therefore biodiversity. For example, kelp holdfasts form intricate structures and host the largest number of organisms in kelp ecosystems. However, methods that quantify 3D habitat complexity have only seldom been used in marine habitats, and never in kelp holdfast communities. This study investigated the role of kelp holdfasts (Laminaria hyperborea) in supporting benthic faunal biodiversity. Computer-aided tomography (CT-) scanning was used to quantify the three-dimensional geometrical complexity of holdfasts, including volume, surface area and surface fractal dimension (FD). Additionally, the number of haptera, number of haptera per unit of volume, and age of kelps were estimated. These measurements were compared to faunal biodiversity and community structure, using partial least-squares regression and multivariate ordination. Holdfast volume explained most of the variance observed in biodiversity indices, however all other complexity measures also strongly contributed to the variance observed. Multivariate ordinations further revealed that surface area and haptera per unit of volume accounted for the patterns observed in faunal community structure. Using 3D image analysis, this study makes a strong contribution to elucidate quantitative mechanisms underlying the observed relationship between biodiversity and habitat complexity. Furthermore, the potential of CT-scanning as an ecological tool is demonstrated, and a methodology for its use in future similar studies is established. Such spatially resolved imager analysis could help identify structurally complex areas as biodiversity hotspots, and may support the prioritization of areas for conservation.

Application of computer-aided tomography techniques to visualize kelp holdfast structure reveals the importance of habitat complexity for supporting marine biodiversity

Ecosystem engineers that increase habitat complexity are keystone species in marine systems, increasing shelter and niche availability, and therefore biodiversity. For example, kelp holdfasts form intricate structures and host the largest number of organisms in kelp ecosystems. However, methods that quantify 3D habitat complexity have only seldom been used in marine habitats, and never in kelp holdfast communities. This study investigated the role of kelp holdfasts (Laminaria hyperborea) in supporting benthic faunal biodiversity. Computer-aided tomography (CT-) scanning was used to quantify the three-dimensional geometrical complexity of holdfasts, including volume, surface area and surface fractal dimension (FD). Additionally, the number of haptera, number of haptera per unit of volume, and age of kelps were estimated. These measurements were compared to faunal biodiversity and community structure, using partial least-squares regression and multivariate ordination. Holdfast volume explained most of the variance observed in biodiversity indices, however all other complexity measures also strongly contributed to the variance observed. Multivariate ordinations further revealed that surface area and haptera per unit of volume accounted for the patterns observed in faunal community structure. Using 3D image analysis, this study makes a strong contribution to elucidate quantitative mechanisms underlying the observed relationship between biodiversity and habitat complexity. Furthermore, the potential of CT-scanning as an ecological tool is demonstrated, and a methodology for its use in future similar studies is established. Such spatially resolved imager analysis could help identify structurally complex areas as biodiversity hotspots, and may support the prioritization of areas for conservation.

Descriptions of Scottish Priority Marine Features (PMFs).

Background The seas around Scotland are rich and diverse – Scotland’s position at the edge of the continental shelf, the long coastline, large area of sea and the mixing of warm and coldwater currents combine to make its waters a special place for marine wildlife and habitats. Scotland has over 18,000 km of coastline and its inshore and offshore areas are among the largest of any EU country, representing 13% of all European seas. Scotland’s seas are of outstanding scenic, historical and cultural value and are part of the national identity at home and abroad. The Marine (Scotland) Act 2010 and the UK Marine and Coastal Access Act 2009 include new powers and duties to ensure that our seas are managed sustainably for future generations, integrating the economic growth of marine industries with the need to protect these assets. Measures to conserve Scotland’s marine natural heritage are based on a three pillar approach, with action at the wider seas level (e.g. marine planning or sectoral controls); specific species conservation measures (e.g. improved protection for seals); and through site protection measures - the identification of new Marine Protected Areas (MPAs). To help target action under each of the three pillars, Scottish Natural Heritage (SNH) and the Joint Nature Conservation Committee (JNCC) have generated a focused list of habitats and species of priority conservation importance - the Priority Marine Features (PMFs). The aim of the current study was to produce a descriptive catalogue of the Scottish PMFs (including component habitats and species where appropriate) to serve as a reference for future nature conservation action. Whilst derived from available existing accounts, the succinct 1-page descriptions are written from a Scottish perspective, refining, but clearly linking to more generic UK, EC or OSPAR (Oslo and Paris Commission) commentary. Available information on the geographic distribution of the features was collated as part of the project and a summary map is provided in each description. Main findings  This project has generated a descriptive catalogue of the 81 PMFs that have been identified in the seas around Scotland (out to the limit of the UK continental shelf). The list comprises 26 broad habitats (e.g. burrowed mud), seven low or limited mobility species (e.g. ocean quahog) and 48 mobile species, including fish (e.g. blue ling) and marine mammals (e.g. minke whale).  Information on the distribution of the PMFs was collated within a Geographic Information System (GIS). This is the first time that data about such a diverse range of Scottish marine nature conservation interests have been compiled within a single repository. These data have and will be used in conjunction with other contextual base-mapping to inform the development of nature conservation advice and commentary (e.g. in the production of the Scotland’s Marine Atlas - Baxter et al., 2011).  The feature distribution mapping used in the production of this report is being made available to view online via the National Marine Plan Interactive web portal (NMPi - http://www.gov.scot/Topics/marine/seamanagement/nmpihome). As new or refined data on Scottish PMFs become available, these will be fed into updates to the project geodatabase and NMPi.

Descriptions of Scottish Priority Marine Features (PMFs).

Background The seas around Scotland are rich and diverse – Scotland’s position at the edge of the continental shelf, the long coastline, large area of sea and the mixing of warm and coldwater currents combine to make its waters a special place for marine wildlife and habitats. Scotland has over 18,000 km of coastline and its inshore and offshore areas are among the largest of any EU country, representing 13% of all European seas. Scotland’s seas are of outstanding scenic, historical and cultural value and are part of the national identity at home and abroad. The Marine (Scotland) Act 2010 and the UK Marine and Coastal Access Act 2009 include new powers and duties to ensure that our seas are managed sustainably for future generations, integrating the economic growth of marine industries with the need to protect these assets. Measures to conserve Scotland’s marine natural heritage are based on a three pillar approach, with action at the wider seas level (e.g. marine planning or sectoral controls); specific species conservation measures (e.g. improved protection for seals); and through site protection measures - the identification of new Marine Protected Areas (MPAs). To help target action under each of the three pillars, Scottish Natural Heritage (SNH) and the Joint Nature Conservation Committee (JNCC) have generated a focused list of habitats and species of priority conservation importance - the Priority Marine Features (PMFs). The aim of the current study was to produce a descriptive catalogue of the Scottish PMFs (including component habitats and species where appropriate) to serve as a reference for future nature conservation action. Whilst derived from available existing accounts, the succinct 1-page descriptions are written from a Scottish perspective, refining, but clearly linking to more generic UK, EC or OSPAR (Oslo and Paris Commission) commentary. Available information on the geographic distribution of the features was collated as part of the project and a summary map is provided in each description. Main findings  This project has generated a descriptive catalogue of the 81 PMFs that have been identified in the seas around Scotland (out to the limit of the UK continental shelf). The list comprises 26 broad habitats (e.g. burrowed mud), seven low or limited mobility species (e.g. ocean quahog) and 48 mobile species, including fish (e.g. blue ling) and marine mammals (e.g. minke whale).  Information on the distribution of the PMFs was collated within a Geographic Information System (GIS). This is the first time that data about such a diverse range of Scottish marine nature conservation interests have been compiled within a single repository. These data have and will be used in conjunction with other contextual base-mapping to inform the development of nature conservation advice and commentary (e.g. in the production of the Scotland’s Marine Atlas - Baxter et al., 2011).  The feature distribution mapping used in the production of this report is being made available to view online via the National Marine Plan Interactive web portal (NMPi - http://www.gov.scot/Topics/marine/seamanagement/nmpihome). As new or refined data on Scottish PMFs become available, these will be fed into updates to the project geodatabase and NMPi.

Acoustic telemetry and network analysis reveal the space use of multiple reef predators and enhance marine protected area design

Marine protected areas (MPAs) are commonly employed to protect ecosystems from threats like overfishing. Ideally, MPA design should incorporate movement data from multiple target species to ensure sufficient habitat is protected. We used long-term acoustic telemetry and network analysis to determine the fine-scale space use of five shark and one turtle species at a remote atoll in the Seychelles, Indian Ocean, and evaluate the efficacy of a proposed MPA. Results revealed strong, species-specific habitat use in both sharks and turtles, with corresponding variation in MPA use. Defining the MPA's boundary from the edge of the reef flat at low tide instead of the beach at high tide (the current best in Seychelles) significantly increased the MPA's coverage of predator movements by an average of 34%. Informed by these results, the larger MPA was adopted by the Seychelles government, demonstrating how telemetry data can improve shark spatial conservation by affecting policy directly.

Acoustic telemetry and network analysis reveal the space use of multiple reef predators and enhance marine protected area design

Marine protected areas (MPAs) are commonly employed to protect ecosystems from threats like overfishing. Ideally, MPA design should incorporate movement data from multiple target species to ensure sufficient habitat is protected. We used long-term acoustic telemetry and network analysis to determine the fine-scale space use of five shark and one turtle species at a remote atoll in the Seychelles, Indian Ocean, and evaluate the efficacy of a proposed MPA. Results revealed strong, species-specific habitat use in both sharks and turtles, with corresponding variation in MPA use. Defining the MPA's boundary from the edge of the reef flat at low tide instead of the beach at high tide (the current best in Seychelles) significantly increased the MPA's coverage of predator movements by an average of 34%. Informed by these results, the larger MPA was adopted by the Seychelles government, demonstrating how telemetry data can improve shark spatial conservation by affecting policy directly.