Intrinsic and extrinsic factors drive ontogeny of early-life at-sea behaviour in a marine top predator

Young animals must learn to forage effectively to survive the transition from parental provisioning to independent feeding. Rapid development of successful foraging strategies is particularly important for capital breeders that do not receive parental guidance after weaning. The intrinsic and extrinsic drivers of variation in ontogeny of foraging are poorly understood for many species. Grey seals (Halichoerus grypus) are typical capital breeders; pups are abandoned on the natal site after a brief suckling phase, and must develop foraging skills without external input. We collected location and dive data from recently-weaned grey seal pups from two regions of the United Kingdom (the North Sea and the Celtic and Irish Seas) using animal-borne telemetry devices during their first months of independence at sea. Dive duration, depth, bottom time, and benthic diving increased over the first 40 days. The shape and magnitude of changes differed between regions. Females consistently had longer bottom times, and in the Celtic and Irish Seas they used shallower water than males. Regional sex differences suggest that extrinsic factors, such as water depth, contribute to behavioural sexual segregation. We recommend that conservation strategies consider movements of young naïve animals in addition to those of adults to account for developmental behavioural changes

A novel approach to using seabed geomorphology as a predictor of habitat use in highly mobile marine predators : implications for ecology and conservation

Department for Business, Energy, and Industrial Strategy (BEIS, formerly DECC) Offshore Energy Strategic Environmental Assessment programme funded the purchase of the tags and their deployment (OESEA-14-47) and supported HMEW (OESEA-20-118). DJFR was supported by NERC National Public Good- National Capability funding to SMRU (NE/R015007/1). MIDC was supported by project EcoSTAR, funded under NERC INSITE Phase II (grant no. NE/T010614/1). Funds for open access publication fees were from the research grants of supervisors at University of St Andrews.Understanding how marine predators find patchily distributed prey resources in a dynamic environment is key to identifying important ecological areas for ecosystem-level conservation management. However, the mechanisms underpinning important foraging areas often result from complex interactions between static and dynamic covariates (e.g. topography and currents). Modelling habitat associations with hydrodynamic processes is rarely useful when attempting to identify and characterise foraging areas across an individual’s foraging range. Investigating the influence of static habitat features on predator behaviour can provide a more tractable baseline understanding of habitat associations, upon which additional complexity can be added. Seabed gradient covariates (e.g. slope and aspect) are often used, yet such metrics are computed at singular user-defined resolutions, and provide limited ecological insight when used in isolation. Instead, categorising the seabed into geomorphological features may provide better characterisation of seabed structure. Here we explore the utility of a pattern recognition algorithm to delineate whole geomorphological features (“geomorphons”) on the seabed (e.g. valleys, ridges, footslopes) from bathymetry data, and examine the influence of geomorphology on marine predator habitat use. We demonstrate the potential application of this approach in a case study, examining the influence of geomorphons on the at-sea behaviour of a highly mobile predator inhabiting shelf seas: the grey seal (Halichoerus grypus). We analyse GPS tracking data from three seals tagged in the southern North Sea, an area with heterogeneous geomorphology. We use hidden Markov models (HMMs) to infer foraging and travelling behaviour and model the effect of different feature types on the probability of switching between states. All three seals showed an increased probability of transitioning from travelling to foraging when encountering slopes, footslopes and hollows, and foraging activity was concentrated at slopes on the fringes of the Dogger Bank. We hypothesise that such features may host prey aggregations, or lead to increased prey capture success. The results suggest the importance of such areas for grey seals in the southern North Sea, a region undergoing rapid and widespread anthropogenic habitat change. This method could be incorporated into future species distribution models to improve estimates of predator distribution, informing conservation management and marine spatial planning.Publisher PDFPeer reviewe

From pup to predator; generalized hidden Markov models reveal rapid development of movement strategies in a naïve long‐lived vertebrate

Rapid development of a successful foraging strategy is critical for juvenile survival, especially for naïve animals that receive no parental guidance. However, this process is poorly understood for many species. Although observation of early‐life movements is increasingly possible with miniaturisation of animal‐borne telemetry devices, analytical limitations remain. Here, we tracked 29 recently‐weaned, grey seal Halichoerus grypus pups from colonies in two geographically distinct regions of the United Kingdom. We analysed at‐sea movements of pups throughout their initial months of nutritional independence to investigate the ontogeny of behaviour‐specific (foraging and travelling) movement patterns. Using generalized hidden Markov models (HMMs), we extended the conventional HMM framework to account for temporal changes in putative foraging and travelling movement characteristics, and investigate the effects of intrinsic (sex) and extrinsic (environment) factors on this process. Putative foraging behaviour became more tortuous with time, and travelling became faster and more directed, suggesting a reduction in search scale and an increase in travel efficiency as pups shifted from exploration to an adult‐like repeatable foraging strategy. Sex differences in movement characteristics were evident from colony departure, but sex‐specific activity budgets were only detected in one region. We show that sex‐specific behavioural strategies emerge before sexual size dimorphism in grey seals, and suggest that this phenomenon may occur in other long‐lived species. Our results also indicate that environmental variation may affect the emergence of sex‐specific foraging behaviour, highlighting the need to consider interacting intrinsic and extrinsic factors in shaping movement strategies of long‐lived vertebrates. Moreover, comparing the behavioural state estimations to those of a conventional HMM (no variation in state‐specific movement parameters) revealed differences in the amount and location of foraging activity, with implications for spatial conservation management. Overlooking intrinsic and extrinsic variation in movement processes could distort our understanding of foraging ecology, population dynamics, and conservation requirements

Sympatric seals, satellite tracking and protected areas : habitat-based distribution estimates for conservation and management

Analysis was funded by the UK Government Department for Business, Energy and Industrial Strategy (BEIS; OESEA-16-76/OESEA-17-78) with support from the Natural Environment Research Council (NERC; INSITE Phase II NE/T010614/1 EcoSTAR), EU INTERREG (MarPAMM), and the Scottish Government (MMSS/002/15). DJFR’s contribution was funded by NERC National Capability Funding (NE/R015007/1). WJG was supported by INSITE Phase I (MAPS). Telemetry tags and their deployment were funded in the UK by BEIS (and previous incarnations), NERC, Marine Scotland, Scottish Government, NatureScot, SMRU, SMRU Instrumentation Group, Marine Current Turbines, Ørsted, the Met Office, the Zoological Society of London (ZSL), the Crown Estate, Highlands & Islands Enterprise, Moray Firth Renewables Limited (MORL), Beatrice Offshore Windfarm Limited (BOWL), SITA Trust, BBC Wildlife Fund and the Hampshire & Isle of Wight Wildlife Trust. Tags and their deployment in Ireland were funded by Inland Fisheries Ireland, the Department of Communications, Marine and Natural Resources, the Higher Education Authority of Ireland, the National Geographic Society, the Department of Agriculture, Food and the Marine, and the National Parks and Wildlife Service. UK aerial surveys conducted by SMRU were funded by NERC (NE/R015007/1), NatureScot, the Department for Agriculture, Environment and Rural Affairs (Northern Ireland), Marine Current Turbines, Marine Scotland, Natural England, and Scottish Power. Aerial surveys in Ireland were funded by the Department for Tourism, Culture, Arts, Gaeltacht, Sport and Media.Marine predator populations are crucial to the structure and functioning of ecosystems. Like many predator taxa, pinnipeds face an increasingly complex array of natural and anthropogenic threats. Understanding the relationship between at-sea processes and trends in abundance at land-based monitoring sites requires robust estimates of at-sea distribution, often on multi-region scales. Such an understanding is critical for effective conservation management, but estimates are often limited in spatial extent by spatial coverage of animal-borne tracking data. Grey (Halichoerus grypus) and harbour seals (Phoca vitulina) are sympatric predators in North Atlantic shelf seas. The United Kingdom (UK) and Ireland represents an important population centre for both species, and Special Areas of Conservation (SACs) are designated for their monitoring and protection. Here we use an extensive high-resolution GPS tracking dataset, unprecedented in both size (114 grey and 239 harbour seals) and spatial coverage, to model habitat preference and generate at-sea distribution estimates for the entire UK and Ireland populations of both species. We found regional differences in environmental drivers of distribution for both species which likely relate to regional variation in diet and population trends. Moreover, we provide SAC-specific estimates of at-sea distribution for use in marine spatial planning, demonstrating that hotspots of at-sea density in UK and Ireland-wide maps cannot always be apportioned to the nearest SAC. We show that for grey seals, colonial capital breeders, there is a mismatch between SACs (where impacts are likely to be detected) and areas where impacts are most likely to occur (at sea). We highlight an urgent need for further research to elucidate the links between at-sea distribution during the foraging season and population trends observed in SACs. More generally, we highlight that the potential for such a disconnect needs to be considered when designating and managing protected sites, particularly for species that aggregate to breed and exhibit partial migration (e.g. grey seals), or spatial variation in migration strategies. We demonstrate the use of strategic tracking efforts to predict distribution across multiple regions, but caution that such efforts should be mindful of the potential for differences in species-environment relationships despite similar accessible habitats.Publisher PDFPeer reviewe

Predicting the exposure of diving grey seals to shipping noise.

There is high spatial overlap between grey seals and shipping traffic, and the functional hearing range of grey seals indicates sensitivity to underwater noise emitted by ships. However, there is still very little data regarding the exposure of grey seals to shipping noise, constraining effective policy decisions. Particularly, there are few predictions that consider the at-sea movement of seals. Consequently, this study aimed to predict the exposure of adult grey seals and pups to shipping noise along a three-dimensional movement track, and assess the influence of shipping characteristics on sound exposure levels. Using ship location data, a ship source model, and the acoustic propagation model, RAMSurf, this study estimated weighted 24-h sound exposure levels (10-1000 Hz) (SELw). Median predicted 24-h SELw was 128 and 142 dB re 1 μPa2s for the pups and adults, respectively. The predicted exposure of seals to shipping noise did not exceed best evidence thresholds for temporary threshold shift. Exposure was mediated by the number of ships, ship source level, the distance between seals and ships, and the at-sea behaviour of the seals. The results can inform regulatory planning related to anthropogenic pressures on seal populations

Gravitating discs around black holes

Fluid discs and tori around black holes are discussed within different approaches and with the emphasis on the role of disc gravity. First reviewed are the prospects of investigating the gravitational field of a black hole--disc system by analytical solutions of stationary, axially symmetric Einstein's equations. Then, more detailed considerations are focused to middle and outer parts of extended disc-like configurations where relativistic effects are small and the Newtonian description is adequate. Within general relativity, only a static case has been analysed in detail. Results are often very inspiring, however, simplifying assumptions must be imposed: ad hoc profiles of the disc density are commonly assumed and the effects of frame-dragging and completely lacking. Astrophysical discs (e.g. accretion discs in active galactic nuclei) typically extend far beyond the relativistic domain and are fairly diluted. However, self-gravity is still essential for their structure and evolution, as well as for their radiation emission and the impact on the environment around. For example, a nuclear star cluster in a galactic centre may bear various imprints of mutual star--disc interactions, which can be recognised in observational properties, such as the relation between the central mass and stellar velocity dispersion.Comment: Accepted for publication in CQG; high-resolution figures will be available from http://www.iop.org/EJ/journal/CQ

Aptamer-based multiplexed proteomic technology for biomarker discovery

Interrogation of the human proteome in a highly multiplexed and efficient manner remains a coveted and challenging goal in biology. We present a new aptamer-based proteomic technology for biomarker discovery capable of simultaneously measuring thousands of proteins from small sample volumes (15 [mu]L of serum or plasma). Our current assay allows us to measure ~800 proteins with very low limits of detection (1 pM average), 7 logs of overall dynamic range, and 5% average coefficient of variation. This technology is enabled by a new generation of aptamers that contain chemically modified nucleotides, which greatly expand the physicochemical diversity of the large randomized nucleic acid libraries from which the aptamers are selected. Proteins in complex matrices such as plasma are measured with a process that transforms a signature of protein concentrations into a corresponding DNA aptamer concentration signature, which is then quantified with a DNA microarray. In essence, our assay takes advantage of the dual nature of aptamers as both folded binding entities with defined shapes and unique sequences recognizable by specific hybridization probes. To demonstrate the utility of our proteomics biomarker discovery technology, we applied it to a clinical study of chronic kidney disease (CKD). We identified two well known CKD biomarkers as well as an additional 58 potential CKD biomarkers. These results demonstrate the potential utility of our technology to discover unique protein signatures characteristic of various disease states. More generally, we describe a versatile and powerful tool that allows large-scale comparison of proteome profiles among discrete populations. This unbiased and highly multiplexed search engine will enable the discovery of novel biomarkers in a manner that is unencumbered by our incomplete knowledge of biology, thereby helping to advance the next generation of evidence-based medicine

Generic thin-shell gravastars

We construct generic spherically symmetric thin-shell gravastars by using the cut-and-paste procedure. We take considerable effort to make the analysis as general and unified as practicable; investigating both the internal physics of the transition layer and its interaction with "external forces" arising due to interactions between the transition layer and the bulk spacetime. Furthermore, we discuss both the dynamic and static situations. In particular, we consider "bounded excursion" dynamical configurations, and probe the stability of static configurations. For gravastars there is always a particularly compelling configuration in which the surface energy density is zero, while surface tension is nonzero.Comment: V1: 39 pages, 9 figures; V2: 40 pages, 9 figures. References added, some discussion added, some typos fixed. Identical to published version. arXiv admin note: text overlap with arXiv:1112.205

Gravitational Lensing by Black Holes

We review the theoretical aspects of gravitational lensing by black holes, and discuss the perspectives for realistic observations. We will first treat lensing by spherically symmetric black holes, in which the formation of infinite sequences of higher order images emerges in the clearest way. We will then consider the effects of the spin of the black hole, with the formation of giant higher order caustics and multiple images. Finally, we will consider the perspectives for observations of black hole lensing, from the detection of secondary images of stellar sources and spots on the accretion disk to the interpretation of iron K-lines and direct imaging of the shadow of the black hole.Comment: Invited article for the GRG special issue on lensing (P. Jetzer, Y. Mellier and V. Perlick Eds.). 31 pages, 12 figure