Bar-headed geese are renowned for migratory flights at extremely high altitudes over the world's tallest mountains, the Himalayas, where partial pressure of oxygen is dramatically reduced while flight costs, in terms of rate of oxygen consumption, are greatly increased. Such a mismatch is paradoxical, and it is not clear why geese might fly higher than is absolutely necessary. In addition, direct empirical measurements of high-altitude flight are lacking. We test whether migrating bar-headed geese actually minimize flight altitude and make use of favourable winds to reduce flight costs. By tracking 91 geese, we show that these birds typically travel through the valleys of the Himalayas and not over the summits. We report maximum flight altitudes of 7290 m and 6540 m for southbound and northbound geese, respectively, but with 95 per cent of locations received from less than 5489 m. Geese travelled along a route that was 112 km longer than the great circle (shortest distance) route, with transit ground speeds suggesting that they rarely profited from tailwinds. Bar-headed geese from these eastern populations generally travel only as high as the terrain beneath them dictates and rarely in profitable winds. Nevertheless, their migration represents an enormous challenge in conditions where humans and other mammals are only able to operate at levels well below their sea-level maxima.

It is now well understood that climate change has the potential to dramatically affect biodiversity, with effects on spatio-temporal distribution patterns, trophic relationships and survivorship. In the marine turtles, sex is determined by incubation temperature, such that warming temperatures could lead to a higher production of female hatchlings. By measuring nest temperature, and using a model to relate the incubation temperature to sex ratio, we estimate that Caribbean Colombian leatherback sea turtles currently produce approximately 92% female hatchlings. We modelled the relationship between incubation, sand and air temperature, and under all future climate change scenarios (0.4-6.0 °C warming over the next 100 years), complete feminization could occur, as soon as the next decade. However, male producing refugia exist in the periphery of smaller nests (0.7 °C cooler at the bottom than at the centre), within beaches (0.3 °C cooler in the vegetation line and inter-tidal zone) and between beaches (0.4 °C higher on dark beaches), and these natural refugia could be assigned preferential conservation status. However, there exists a need to develop strategies that may ameliorate deleterious effects of climate-induced temperature changes in the future. We experimentally shaded clutches using screening material, and found that it was effective in reducing nest temperature, producing a higher proportion of male hatchlings, without compromising the fitness or hatching success. Artificial shade in hatcheries is a very useful and simple tool in years or periods of high environmental temperatures. Nevertheless, this is only an emergency response to the severe impacts that will eventually have to be reversed if we are to guarantee the stability of the populations. © 2011 Blackwell Publishing Ltd.

Conserving and managing populations of marine vertebrates can be complex when they occupy the waters of multiple nations, crossing heterogeneous legal and management landscapes. Hawksbill turtles (. Eretmochelys imbricata) are subject to varying levels of use in Caribbean countries and their conservation and management is complicated by the extent to which they are a 'shared' resource. In 1997 and 2000, Cuba attempted to 'downlist' hawksbills from Cuban waters to CITES Appendix II to allow limited international trade. The research on movement and dispersal of hawksbills reported here was undertaken to better inform discussion about the impacts of their harvest. Flipper tagging and satellite tracking demonstrate that the majority of study turtles remained in Cuban territorial waters. of 1170 hawksbills tagged (525 adults and 606 juveniles), 12% (. n=. 143) were recaptured. All recaptured adults (. n=. 16 males, 38 nesting females and 30 adult females in-water) were in Cuban waters. of the 59 juveniles recaptured, only four recaptures were outside Cuban waters (Nicaragua. =. 2, Colombia. =. 1, USA. =. 1). Fourteen hawksbills tagged in the waters of other nations were recaptured in Cuban waters. We also satellite tracked 21 turtles (one adult male, ten nesting females and ten non-nesting adult females), of which five tags failed, 11 stayed in Cuban waters for the duration of transmissions (1-809. days) and five foraged in the waters of other nations (Mexico. n=. 1 an adult female; Honduras. n=. 2, both post-nesting turtles; Colombia. n=. 1 an adult female; and the eastern Lesser Antilles. n=. 1, an adult male), with differences for nesting and non-nesting turtles. Our results, demonstrating extended site fidelity within Cuban waters, suggest that strengthening national management within national jurisdictions that host hawksbill turtles is fundamental to improving regional conservation as a whole. © 2012 Elsevier Ltd.

Birds that fly over mountain barriers must be capable of meeting the increased energetic cost of climbing in low-density air, even though less oxygen may be available to support their metabolism. This challenge is magnified by the reduction in maximum sustained climbing rates in large birds. Bar-headed geese (Anser indicus) make one of the highest and most iconic transmountain migrations in the world. We show that those populations of geese that winter at sea level in India are capable of passing over the Himalayas in 1 d, typically climbing between 4,000 and 6,000 m in 7-8 h. Surprisingly, these birds do not rely on the assistance of upslope tailwinds that usually occur during the day and can support minimum climb rates of 0.8-2.2 km·h(-1), even in the relative stillness of the night. They appear to strategically avoid higher speed winds during the afternoon, thus maximizing safety and control during flight. It would seem, therefore, that bar-headed geese are capable of sustained climbing flight over the passes of the Himalaya under their own aerobic power.

Background: Hawksbill turtles (Eretmochelys imbricata) are Critically Endangered throughout their global range, and concerningly little is known about this species in the Red Sea. With large-scale coastal development projects underway in the northern Red Sea, it is critical to understand the movement and habitat use patterns of hawksbill turtles in this environmentally unique region, so that effective conservation strategies can be implemented. We satellite tagged three hawksbill turtles, one 63 cm curved carapace length adult male captured near Wahlei Island, one 55 cm turtle captured in the Gulf of Aqaba, and one 56 cm turtle suffering from a floating syndrome which was captured at Waqqadi Island, rehabilitated, and released at Waqqadi Island. Turtles were tracked for 156, 199, and 372 days between October 2020 and November 2021. Results: We calculated the home ranges and core use areas of hawksbill turtles using kernel-density estimations and found that each turtle showed high fidelity to their foraging sites. Home ranges calculated with GPS-derived locations ranged between 13.6 and 2.86 km2, whereas home ranges calculated with Argos-derived locations ranged from 38.98 to 286.45 km2. GPS-derived locations also revealed a higher proportion of time spent in coral and rock habitats compared to Argos, based on location overlap with the Allen Coral Reef Atlas. We also found that turtles were making shallow dives, usually remaining between 0 and 5 m. Conclusions: While the number of tracked turtles in this study was small, it represents an important contribution to the current understanding of spatial ecology among foraging hawksbill turtles globally, and provides the first-ever reported hawksbill turtle tracking data from the Red Sea. Our results suggest that protecting coral reef habitats and implementing boating speed limits near reefs could be effective conservation measures for foraging hawksbill turtles in the face of rapid coastal development.

The Red Sea is particularly biodiverse, hosting high levels of endemism and numerous populations whose extinction risk is heightened by their relative isolation. Elasmobranchs and sea turtles have likely suffered recent declines in this region, although data on their distribution and biology are severely lacking, especially on the eastern side of the basin in Saudi Arabian waters. Here, we present sightings of elasmobranchs and sea turtles across the north-eastern Red Sea and Gulf of Aqaba collected through a combination of survey methods. Over 455 survey hours, we recorded 407 sightings belonging to 26 elasmobranch species and two sea turtle species, more than 75% of which are of conservation concern. We identified 4 species of rays and 9 species of sharks not previously recorded in Saudi Arabia and report a range extension for the pink whipray (Himantura fai) and the round ribbontail ray (Taeniurops meyeni) into the Gulf of Aqaba. High density of sightings of conservation significance, including green and hawksbill sea turtles and halavi guitarfish were recorded in bay systems along the eastern Gulf of Aqaba and the Saudi Arabian coastline bordering the north-eastern Red Sea, and many carcharhinid species were encountered at offshore seamounts in the region. Our findings provide new insights into the distribution patterns of megafaunal assemblages over smaller spatial scales in the region, and facilitate future research and conservation efforts, amidst ongoing, large-scale coastal developments in the north-eastern Red Sea and Gulf of Aqaba.

Knowledge of the three-dimensional movement patterns of elasmobranchs is vital to understand their ecological roles and exposure to anthropogenic pressures. To date, comparative studies among species at global scales have mostly focused on horizontal movements. Our study addresses the knowledge gap of vertical movements by compiling the first global synthesis of vertical habitat use by elasmobranchs from data obtained by deployment of 989 biotelemetry tags on 38 elasmobranch species. Elasmobranchs displayed high intra- and interspecific variability in vertical movement patterns. Substantial vertical overlap was observed for many epipelagic elasmobranchs, indicating an increased likelihood to display spatial overlap, biologically interact, and share similar risk to anthropogenic threats that vary on a vertical gradient. We highlight the critical next steps toward incorporating vertical movement into global management and monitoring strategies for elasmobranchs, emphasizing the need to address geographic and taxonomic biases in deployments and to concurrently consider both horizontal and vertical movements.

During the breeding season, seabirds are central place foragers and in order to successfully rear chicks they must adjust their foraging behaviours to compensate for extrinsic factors. When foraging, arctic terns Sterna paradisaea are restricted to the first 50 cm of the water column and can only carry a few prey items back to their nests at once. In Iceland, where 20–30% of the global population breed, poor fledging success has been linked to low food availability. Using GPS loggers, we investigated individual foraging behaviours of breeding adults during incubation from a large colony over four seasons. First, we tested whether foraging trip distance or duration was linked to morphology or sex. Second, we examined how trips vary with weather and overlap with commercial fisheries. Our findings reveal that arctic terns travel far greater distances during foraging trips than previously recorded (approximately 7.3 times further), and they forage around the clock. There was inter-annual variability in the foraging locations that birds used, but no relationship between size or sex differences and the distances travelled. We detected no relationship between arctic tern foraging flights and local prevailing winds, and tern heading and speed were unrelated to local wind patterns. We identified key arctic tern foraging areas and found little spatial or temporal overlap with fishing pelagic vessels, but larger home ranges corresponded with years with lower net primary productivity levels. This suggests that whilst changing polar weather conditions may not pose a threat to arctic terns at present, nor might local competition with commercial fisheries for prey, they may be failing to forage in productive areas, or may be affected by synergistic climatic effects on prey abundance and quality. Shifts in pelagic prey distributions as a result of increasing water temperatures and salinities will impact marine top predators in this region, so continued monitoring of sentinel species such as arctic terns is vital.

Wildlife rescue and rehabilitation is used globally to aid the conservation and welfare of marine species, however, postrelease monitoring is challenging. Here, long-term, regional postrelease monitoring provides feedback for rehabilitation centers for gray seals (Halichoerus grypus). Data from 1,094 rehabilitated gray seals over 19 years across the southwest UK were examined to assess postrelease survivorship and the impact of release site on movements and range. Using flipper tags combined with photo identification, 391 rehabilitated seals (35.7%) were resighted, including 188 seals (17.2%) that were traced back to a specific rehabilitated individual with release data. The maximum monitoring duration for a single rehabilitated seal was 17 years, although the majority (151/188; 80%) were sighted for less than 5 years and 80/188 (43%) were resighted for less than a year. Almost all 188 traced rehabilitated seals (n = 176, 93.6%) visited the St Ives Bay Wild Site, yet only half had been released at the adjacent St Ives Bay Release Site. Rehabilitated seals had similar dispersal patterns to their wild conspecifics but over a smaller area. Once released, rehabilitated animals face the same threats as their wild counterparts.

Climate change is a threat to marine turtles that is expected to affect all of their life stages. To guide future research, we conducted a review of the most recent literature on this topic, highlighting knowledge gains and research gaps since a similar previous review in 2009. Most research has been focussed on the terrestrial life history phase, where expected impacts will range from habitat loss and decreased reproductive success to feminization of populations, but changes in reproductive periodicity, shifts in latitudinal ranges, and changes in foraging success are all expected in the marine life history phase. Models have been proposed to improve estimates of primary sex ratios, while technological advances promise a better understanding of how climate can influence different life stages and habitats. We suggest a number of research priorities for an improved understanding of how climate change may impact marine turtles, including: improved estimates of primary sex ratios, assessments of the implications of female-biased sex ratios and reduced male production, assessments of the variability in upper thermal limits of clutches, models of beach sediment movement under sea level rise, and assessments of impacts on foraging grounds. Lastly, we suggest that it is not yet possible to recommend manipulating aspects of turtle nesting ecology, as the evidence base with which to understand the results of such interventions is not robust enough, but that strategies for mitigation of stressors should be helpful, providing they consider the synergistic effects of climate change and other anthropogenic-induced threats to marine turtles, and focus on increasing resilience.

Thus far, ecophysiology research has predominantly been conducted within controlled laboratory-based environments, owing to a mismatch between the recording technologies available for physiological monitoring in wild animals and the suite of behaviours and environments they need to withstand, without unduly affecting subjects. While it is possible to record some physiological variables for free-living animals using animal-attached logging devices, including inertial-measurement, heart-rate and temperature loggers, the field is still in its infancy. In this opinion piece, we review the most important future research directions for advancing the field of ‘physiologging’ in wild animals, including the technological development that we anticipate will be required, and the fiscal and ethical challenges that must be overcome. Non-invasive, multi-sensor miniature devices are ubiquitous in the world of human health and fitness monitoring, creating invaluable opportunities for animal and human physiologging to drive synergistic advances. We argue that by capitalizing on the research efforts and advancements made in the development of human wearables, it will be possible to design the non-invasive loggers needed by ecophysiologists to collect accurate physiological data from free-ranging animals ethically and with an absolute minimum of impact. In turn, findings have the capacity to foster transformative advances in human health monitoring. Thus, we invite biomedical engineers and researchers to collaborate with the animal-tagging community to drive forward the advancements necessary to realize the full potential of both fields.
. This article is part of the theme issue ‘Measuring physiology in free-living animals (Part II)’.

The study presents the first national assessment of a nurse shark (Ginglymostoma cirratum) population, conducted using a combination of transect surveys and baited remote underwater videos (BRUVs). Density of nurse sharks in Belize was found to be higher in reefs than in lagoons, and in the atolls furthest away from the mainland and human settlements. Only large and old protected areas were found to have a positive impact on nurse shark abundance. Absolute abundance of nurse sharks was estimated using distance sampling analysis, giving a total nurse shark population in the range of 3,858 to 14,375 sharks. Thanks to a vast area of suitable habitat for nurse sharks in the country and legislation already in place for the safeguard of the species, Belize could represent an important hotspot for nurse sharks in the Western Atlantic. The data presented here hence offers a baseline for the long-term monitoring of the Belizean nurse shark population and improves our understanding of nurse shark abundance and distribution in the wider Caribbean basin.

Environmental change and biodiversity loss are but two of the complex challenges facing conservation practitioners and policy makers. Relevant and robust scientific knowledge is critical for providing decision-makers with the actionable evidence needed to inform conservation decisions. In the Anthropocene, science that leads to meaningful improvements in biodiversity conservation, restoration and management is desperately needed. Conservation Physiology has emerged as a discipline that is well-positioned to identify the mechanisms underpinning population declines, predict responses to environmental change and test different in situ and ex situ conservation interventions for diverse taxa and ecosystems. Here we present a consensus list of 10 priority research themes. Within each theme we identify specific research questions (100 in total), answers to which will address conservation problems and should improve the management of biological resources. The themes frame a set of research questions related to the following: (i) adaptation and phenotypic plasticity; (ii) human-induced environmental change; (iii) human-wildlife interactions; (iv) invasive species; (v) methods, biomarkers and monitoring; (vi) policy, engagement and communication; (vii) pollution; (viii) restoration actions; (ix) threatened species; and (x) urban systems. The themes and questions will hopefully guide and inspire researchers while also helping to demonstrate to practitioners and policy makers the many ways in which physiology can help to support their decisions.

The goal of achieving enhanced diagnosis and continuous monitoring of human health has led to a vibrant, dynamic and well-funded field of research in medical sensing and biosensor technologies. The field has many sub-disciplines which focus on different aspects of sensor science; engaging engineers, chemists, biochemists and clinicians, often in interdisciplinary teams. The trends which dominate include the efforts to develop effective point of care tests and implantable/wearable technologies for early diagnosis and continuous monitoring. This review will outline the current state of the art in a number of relevant fields, including device engineering, chemistry, nanoscience and biomolecular detection, and suggest how these advances might be employed to develop effective systems for measuring physiology, detecting infection and monitoring biomarker status in wild animals. Special consideration is also given to the emerging threat of antimicrobial resistance and in the light of the current SARS-CoV-2 outbreak, zoonotic infections. Both of these areas involve significant crossover between animal and human health and are therefore well placed to seed technological developments with applicability to both human and animal health and, more generally, the reviewed technologies have significant potential to find use in the measurement of physiology in wild animals.This article is part of the theme issue ‘Measuring physiology in free-living animals (Part II)’.

This thesis advances the understanding of the spatial and behavioural ecology of two endangered species, basking sharks (Cetorhinus maximus) and green turtles (Chelonia mydas) at their coastal foraging sites while highlighting the pragmatic application of biologging technology in informing conservation. Chapter 1 of this thesis is a general introduction to tracking technologies, covering how the advancements of biologging have revolutionised the field of ecology, with an emphasis on how accelerometers can be used in conservation. Chapters 2 explores the use of accelerometers on three basking sharks in the western Scottish Isles to understand their fine-scale behaviour. I present early evidence of potential behavioural lateralisation, and the first direct records of 67 breaches over 41 days, with sharks breaching on average twice a day, both during night and day and increasing energy expenditure by at least 30 times to breach. While the function of breaching remains unclear, owing to its energetic cost, breaching is likely to have an important fitness function. In Chapter 3, accelerometers were deployed on 16 juvenile green turtles in the Bahamas to investigate the behavioural and energetic costs of translocation. Turtles rehomed in as little as 15 hours following translocation of 4 km and allocated twice as much time to energetically demanding behaviours compared to resident turtles at their foraging grounds, highlighting that translocation is not a suitable conservation practice for sea turtles. Chapter 4 summarises both data chapters and discusses how their findings provide further evidence of how tracking technologies can be ideal tools for conservation practitioners by monitoring animal movement, behaviour and health as well as assisting with the designation of protected areas by identifying important life history events. Chapter 4 concludes on the challenges and limitations of the thesis as well as the future directions in the use of tracking technologies in conservation ecology.

The physiological mechanisms by which animals regulate energy expenditure, respond to stimuli and stressors, and maintain homeostasis at the tissue, organ and whole organism levels can be described by ‘physiologging’—that is, the use of onboard miniature electronic devices to record physiological metrics of animals in captivity or free-living in the wild. Despite its origins in the 1960s, physiologging has evolved more slowly than its umbrella field of biologging. However, the recording of physiological metrics in free-living animals will be key to solving some of the greatest challenges in biodiversity conservation, issues pertaining to animal health and welfare, and for inspiring future therapeutic strategies for human health. Current physiologging technologies encompass the measurement of physiological variables such as heart rate, brain activity, body temperature, muscle stimulation and dynamic movement, yet future developments will allow for onboard logging of metrics relating to organelle, molecular and genetic function.
. This article is part of the theme issue ‘Measuring physiology in free-living animals (Part II)’.

The Intergovernmental Panel on Climate Change predicts that sea levels will rise by up to 0.82 m in the next 100 years. In natural systems, coastlines would migrate landwards, but because most of the world’s human population occupies the coast, anthropogenic structures (such as sea walls or buildings) have been constructed to defend the shore and prevent loss of property. This can result in a net reduction in beach area, a phenomenon known as “coastal squeeze”, which will reduce beach availability for species such as marine turtles. As of yet, no global assessment of potential future coastal squeeze risk at marine turtle nesting beaches has been conducted. We used Google Earth satellite imagery to enumerate the proportion of beaches over the global nesting range of marine turtles that are backed by hard anthropogenic coastal development (HACD). Mediterranean and North American nesting beaches had the most HACD, while the Australian and African beaches had the least. Loggerhead and Kemp’s ridley turtle nesting beaches had the most HACD, and flatback and green turtles the least. Future management approaches should prioritise the conservation of beaches with low HACD to mitigate future coastal squeeze.

High-altitude environments present challenges to animal life, including cold temperatures, dehydration and reduced barometric pressure, resulting in diminished oxygen availability (hypoxia). Birds that migrate across high-altitude mountain ranges face these challenges whilst meeting the high metabolic demands of flight. In this thesis I develop scientific understanding of how avian migrants overcome this challenge and cross the Tibetan Plateau (the highest and largest land mass on Earth). Firstly, I consolidate research into the benefits that avian physiology can confer for funding high rates of metabolism at high altitude, relative to humans and other mammals. Secondly, I present empirical work that aims to investigate high-altitude flight during migration across the Tibetan Plateau, expanding the scientific literature that has previously focused on bar-headed geese (Anser indicus). I collate tracking data for nine species of birds that migrate across or around the Himalayas and Tibetan Plateau and investigate whether there are common techniques that reduce flight costs and minimise exposure to hypoxia. I also explore how thermoregulation in bar-headed geese may be affected by high altitude, using high resolution data collected during free flight over the Tibetan Plateau. Finally, I compare the muscular phenotype of high-altitude migrant ruddy shelduck (Tadorna ferruginea) and low-altitude congener, common shelduck (Tadorna tadorna), to investigate whether ruddy shelduck have physiological adaptations for flight at high altitude. The research carried out in this thesis reveals that i) multiple species reach altitudes in excess of 6,000 m whilst crossing the Tibetan Plateau, including cranes, raptors and waterfowl who’s flight altitudes have not been previously quantified; ii) with the exception of demoiselle cranes, most bird species reduce altitude exposure where possible; iii) bar-headed geese maintain core body temperatures within a relatively narrow range regardless of altitude, expanding our understanding of the high-altitude physiology of this iconic species; iv) ruddy shelduck may have adaptations to enhance the functional properties of mitochondrial enzymes, which may support high-altitude flight. Collectively, this work advances our knowledge of how migrants overcome the challenge of migration across high altitude through both physiological and behavioural adaptation and adjustment.

The establishment of the piscivorous lionfishPteroisspp. in the Western Atlantic and wider Caribbean is a well-documented example of a successful marine invasion. Recently, lionfish have been shown to colonise a wide range of ecosystems and tolerate a wider range of salinities than previously thought. In the present study, lionfish were maintained in aquaria under differing salinity treatments (10, 20 and 37 psu) similar to those they might experience in an estuarine ecosystem. The effects of long-term hyposaline exposure on growth, metabolic rate, maximum food consumption and digestion were examined. Consistent with previous studies, lionfish were able to survive in hyposaline conditions for extended periods of time. However, lionfish in the most hyposaline treatment (10 psu) exhibited reduced growth under low food conditions, lower maximum metabolic rate, lower aerobic scope, lower maximum food consumption, took longer to digest a standardized meal size and occupied a greater percentage of their aerobic scope during digestion. Results suggest that (1) given the ability of lionfish to tolerate low salinity, updated range expansion models should incorporate salinity data to improve accuracy of predicted range expansion and (2) the invasion of lionfish into low salinity ecosystems, although a serious concern, will not likely lead to the same level of population increase observed for coral reef habitats due to the physiological costs associated with living in low salinities.

This thesis brings together multiple strands of information at different temporal and spatial scales to shed new light on foraging behaviour. Foraging accounts for much of the time and energy budgets of wild animals and underpins many ecological and evolutionary processes. I used the central place foraging trips of a wide-ranging marine predator, the northern gannet Morus bassanus, as a model to study foraging ecology and the influences of fisheries and climate change. My five data chapters span a range of temporal scales, from two-second behavioural classification to interannual variation across 11 years, as well as a range of spatial scales from individual bird-boat interactions to a multi-colony comparison spanning 83% of the latitudinal range of breeding colonies. First, I found sex-specific foraging behaviour across a range of spatial, temporal and isotopic axes and that the extent of segregation varied inter-annually. Second, I examined the energetics of foraging behaviour, revealing the relative costs of different behaviours and how they relate to three-dimensional flight patterns. I found that effort was driven by the time spent performing behaviours rather than differences in the costs of behaviours per unit time. Furthermore, the proportions of each trip spent performing each behaviour were independent of trip length, meaning that foraging trip duration was a good metric of overall effort. Third, I used bird-borne videos cameras to reveal a high incidence of gannets scavenging from fisheries discards and trawl nets in the Celtic Sea. Scavenging is often thought to be an energy-saving strategy, but the energetic cost of scavenging was no different from that of natural foraging. Fourth, I found that gannets did not respond to nearby fishing vessels in Iceland, where discarding is banned and foraging effort suggested plentiful natural prey. Finally, I collated a large multi-colony dataset, which showed that foraging range and trip durations decreased with latitude when controlling for colony size. I related this pattern to poleward range shifting in response to climate change. Overall, I found that foraging behaviour is highly variable in some respects and consistent in others. This is driven by stable factors such as sex and colony location and changing factors such as population size, climate and oceanography, and fisheries practice or policy. This thesis particularly highlights the benefits of combining data from multiple sources, time periods and regions in order to reveal new patterns.

Applying physiological tools, knowledge and concepts to understand conservation problems (i.e. conservation physiology) has becomecommonplace and confers an ability to understand mechanistic processes,develop predictive models and identify cause-and-effect relationships. Conservation physiology is making contributions to conservation solutions; the number of 'success stories' is growing, but there remain unexplored opportunities for which conservation physiology shows immense promise and has the potential to contribute to major advances in protecting and restoring biodiversity. Here, we consider howconservation physiology has evolved with a focus on reframing the discipline to be more inclusive and integrative.Using a 'horizon scan',we further exploreways in which conservation physiology can be more relevant to pressing conservation issues of today (e.g. addressing the Sustainable Development Goals; delivering science to support the UN Decade on Ecosystem Restoration), aswell as more forward-looking to inform emerging issues and policies for tomorrow. Our horizon scan provides evidence that, as the discipline of conservation physiology continues to mature, it provides a wealth of opportunities to promote integration, inclusivity and forward-thinking goals that contribute to achieving conservation gains. To advance environmentalmanagementand ecosystemrestoration,we need to ensure that the underlying science (such as that generated by conservation physiology) is relevant with accompanying messaging that is straightforward and accessible to end users.

AbstractMigratory movements in response to seasonal resources often influence population structure and dynamics. Yet in mobile marine predators, population genetic consequences of such repetitious behaviour remain inaccessible without comprehensive sampling strategies. Temporal genetic sampling of seasonally recurring aggregations of planktivorous basking sharks, Cetorhinus maximus, in the Northeast Atlantic (NEA) affords an opportunity to resolve individual re-encounters at key sites with population connectivity and patterns of relatedness. Genetic tagging (19 microsatellites) revealed 18% of re-sampled individuals in the NEA demonstrated inter/multi-annual site-specific re-encounters. High genetic connectivity and migration between aggregation sites indicate the Irish Sea as an important movement corridor, with a contemporary effective population estimate (Ne) of 382 (CI = 241–830). We contrast the prevailing view of high gene flow across oceanic regions with evidence of population structure within the NEA, with early-season sharks off southwest Ireland possibly representing genetically distinct migrants. Finally, we found basking sharks surfacing together in the NEA are on average more related than expected by chance, suggesting a genetic consequence of, or a potential mechanism maintaining, site-specific re-encounters. Long-term temporal genetic monitoring is paramount in determining future viability of cosmopolitan marine species, identifying genetic units for conservation management, and for understanding aggregation structure and dynamics.

Few studies have looked into climate change resilience of populations of wild animals. We use a model higher vertebrate, the green sea turtle, as its life history is fundamentally affected by climatic conditions, including temperature-dependent sex determination and obligate use of beaches subject to sea level rise (SLR). We use empirical data from a globally important population in West Africa to assess resistance to climate change within a quantitative framework. We project 200 years of primary sex ratios (1900-2100) and create a digital elevation model of the nesting beach to estimate impacts of projected SLR. Primary sex ratio is currently almost balanced, with 52% of hatchlings produced being female. Under IPCC models, we predict: (a) an increase in the proportion of females by 2100 to 76%-93%, but cooler temperatures, both at the end of the nesting season and in shaded areas, will guarantee male hatchling production; (b) IPCC SLR scenarios will lead to 33.4%-43.0% loss of the current nesting area; (c) climate change will contribute to population growth through population feminization, with 32%-64% more nesting females expected by 2120; (d) as incubation temperatures approach lethal levels, however, the population will cease growing and start to decline. Taken together with other factors (degree of foraging plasticity, rookery size and trajectory, and prevailing threats), this nesting population should resist climate change until 2100, and the availability of spatial and temporal microrefugia indicates potential for resilience to predicted impacts, through the evolution of nest site selection or changes in nesting phenology. This represents the most comprehensive assessment to date of climate change resilience of a marine reptile using the most up-to-date IPCC models, appraising the impacts of temperature and SLR, integrated with additional ecological and demographic parameters. We suggest this as a framework for other populations, species and taxa.

The study of animal diet has long been a fundamental area of biological sciences and has developed significantly over the centuries. An understanding of animal diet goes beyond species-specific biology providing insights into interspecific interactions and whole ecosystem functions essential for ecosystem-based management. Anthropogenic activities have caused major declines in marine vertebrate populations many of which are top predators. It is therefore vital to establish an ecological understanding of mesopredators in the oceans who may either mediate or exacerbate the cascading impacts of such declines. Elasmobranch batoids (otherwise referred to as rays) are a diverse, yet highly vulnerable group of mesopredators many of which are considered data deficient and lack comprehensive dietary assessments. Studies that do exist use stomach content analysis (SCA) or stable isotope analysis (SIA) independently of one another. In contrast, the present thesis integrates these two methods. Chapter one aimed to utilize both SCA and SIA techniques to provide the first integrative dietary assessment of the southern stingray (Hypanus americanus), and the first ever quantitative dietary study of the Caribbean whiptail ray (Styracura schmardae), two sympatric and data deficient species. Our results suggested that the diets of both species were similar in structure and composition, though Caribbean whiptail ray diet was dominated by arthropod and annelid prey, while southern stingray diet was dominated by molluscs. The broad variety of taxa identified in the stomachs of both species indicates opportunistic feeding, likely as mesopredators at a trophic level similar to other ray species in their respective families. Our integration of SCA and SIA highlights the advantages of combining the two methods, for example, the higher representation of soft-bodied prey in stable isotope mixed models (SIMM) compared to those of SCA. The focus of Chapter two was the isotopic variances between three metabolically different tissues (blood, white muscle and barb) from both the southern stingray and the Caribbean whiptail ray, highlighting how the use of multiple tissues in diet assessments may give better insight into the temporal variability of diet. This was the first quantitative comparison of SCA and SIA across tissue types in rays, the results of which suggest that method agreement is influenced by tissue type incorporated in SIMM. A limitation of our inferences, however, is the lack of data available on isotopic turnover rates in ray and elasmobranch tissues, thus we recognise the need for further literature and diet manipulation experiments. Nonetheless, the results of the present thesis provide a novel insight into the diets of these two data deficient stingray species and highlight potential avenues for future research which would improve our understanding of these ecologically significant and vulnerable animals.

In this thesis I use stable isotope analysis to investigate the spatial and dietary ecology of two species of tropical stingray, the southern stingray (Hypanus americanus) and the Caribbean whiptail ray (Styracura schmardae) from Eleuthera island, the Bahamas. In Chapter 1, I directly compare stable isotopes of carbon, nitrogen and sulphur between the two species (S. schmardae, n = 96 ; H. americanus, n = 102) to investigate if and how these sympatric stingrays exhibit resource partitioning. I show that mangrove creek systems may be important habitat for S. schmardae, mitigating competition with H. americanus, and that trophic resource partitioning may also be occurring, with H. americanus feeding at a higher trophic level than S. schmardae. In Chapter 2, I explore the use of stable isotope analysis in detecting ontogenetic shifts in H. americanus (n = 110) and S. schmardae (n = 94). Here, I use breakpoint analysis to pinpoint shifts in mean δ15N and δ13C as body size increases, on three metabolically distinct tissues, which therefore give insights into different time periods: whole blood, white muscle and cartilage (barb). There were four breakpoints in white muscle samples, two in blood and in cartilage only one. We recommend that future research determining ontogenetic shifts via stable isotopes utilise this range of tissues. Breakpoints in δ13C were observed in both species, indicating ontogenetic habitat shifts occurring at juvenile sizes. A second shift was detected at larger body sizes in both δ15N and δ13C for S. schmardae, we suggest this second ontogenetic niche shift indicates a return to mangroves and concurrent increase in higher trophic level prey by adults. The findings presented in this thesis are novel for both species, emphasising the significance of mangroves habitats as well as providing the first ever assessment of resource use by the poorly studied Caribbean whiptail ray. Findings could be used to build conservation frameworks to protect southern stingrays, Caribbean whiptail rays, and the mangroves that appear to be intrinsic to their ecology.

In this thesis, I use physiological and behavioural experiments in a laboratory environment to assess the impacts and potential effects of the invasive lionfish (Pterois spp. (Wilcox et al. 2018)) in a mangrove ecosystem. Chapter 1 of the thesis is a general introduction to the main theme of the thesis “lionfish in mangroves”. In this chapter, I present an introduction to invasive species, a general background about the introduction of lionfish in the Western Atlantic and wider Caribbean, the importance of mangrove ecosystems, and then end with the recent discovery of lionfish in mangrove ecosystems. Chapters 2 and 3 are the data chapters of the thesis. In Chapter 2, lionfish were reared in aquaria in the Bahamas under differing salinity treatments (37 ppt (typical reef ecosystem salinity), 20 ppt (typical mangrove ecosystem salinity) and 10 ppt (typical mangrove ecosystem salinity during low tide)) for 84 days. This experiment investigated the effect of acute hyposalinity on growth rate, metabolic rate (standard metabolic rate (SMR), maximal metabolic rate (MMR) and aerobic scope (AS)), maximum food consumption, feeding rates and specific dynamic action (SDAMax, SDAScope, SDATotal, SDADuration). Most lionfish survived in hyposaline conditions for 84 days until the conclusion of the experiment, however, various aspects of their physiology were effected. Lionfish feeding and aerobic capacity (MMR and AS) were reduced, whereas digestive duration (SDADuration) increased at the lowest salinity. Therefore, I suggest: 1) given the ability of lionfish to tolerate low salinity, updated range expansion models should incorporate salinity data to improve accuracy of predicted range expansion for lionfish, and 2) the invasion of lionfish in mangrove habitats with low salinity, although a serious concern, will unlikely lead to the same level of population increase, habitat destruction and decline in native fish populations as observed for some Caribbean coral reef habitats. In Chapter 3, I explore the potential impact of lionfish on a native fish species, which typically occupies a similar trophic niche as lionfish in an invaded mangrove ecosystem, the Schoolmaster snapper (Lutjanus apodus). Using a habitat competition experiment in the laboratory at two salinity treatments (10 ppt (typical mangrove ecosystem salinity during low tide) and 37 ppt (typical reef ecosystem salinity)), I investigated the behavioural interactions between invasive lionfish and native Schoolmaster snapper in a simulated mangrove ecosystem. Schoolmaster snapper showed, on average, a 53% reduction in shelter use when in the presence of lionfish. Results suggest that lionfish have the ability to displace snapper from shelter and are therefore likely to increase the vulnerability of native mangrove fish species to predation by other mangrove meso-predators. Using a behavioural ethogram, I further identified key behaviours displayed by both fish species during a period in which lionfish and snapper both had access to shelter. However, salinity had no effect on the occurrence and outcome of these behaviours, despite my findings in Chapter 2. Chapter 4 is the general discussion of the thesis where I draw on previous research, as well as research from this study to place in context the novel threat of the lionfish invasion in mangroves.
Overall, this thesis investigates the invasiveness of lionfish in a mangrove ecosystem by demonstrating their ability to tolerate and function in low salinity water for extended periods.

Grey seals (Halichoerus grypus) of the North-east Atlantic are protected at designated European Marine Sites (Special Areas of Conservation, SACs) typically during their reproductive periods and in the UK at Sites of Special Scientific Interest (SSSI). As a mobile marine species, grey seals spend other parts of their annual life cycle in non-designated habitat. There is limited information on individual grey seal movements in south-west England. Citizen science photo identification (PID) revealed the movements of 477 grey seals at a regional scale (54 haul-outs up to 230 km apart) for over a decade. Reconstructed movements showed considerable individual variability. Four SACs were linked to up to 18 non-designated sites and two SSSIs in Cornwall were linked to a maximum of 41 non-designated sites. Observations support the value of existing SSSIs at both the well-connected West and North Cornwall sites. Thirteen Marine Protected Areas (MPAs) were visited by grey seals from four SACs and two SSSIs in Cornwall. As a mobile species, grey seals could be included in English MPA management plans. The application of functional linkage from SACs and SSSIs, informed by the movements evidenced in this research, could aid management efforts. This analysis reveals grey seal movements occur across a complex network of interconnected designated and non-designated sites that need to be managed holistically for this species for which the UK has a special responsibility.

The bar-headed goose is famed for migratory flight at extreme altitude. To better understand the physiology underlying this remarkable behavior, we imprinted and trained geese, collecting the first cardiorespiratory measurements of bar-headed geese flying at simulated altitude in a wind tunnel. Metabolic rate during flight increased 16-fold from rest, supported by an increase in the estimated amount of O2 transported per heartbeat and a modest increase in heart rate. The geese appear to have ample cardiac reserves, as heart rate during hypoxic flights was not higher than in normoxic flights. We conclude that flight in hypoxia is largely achieved via the reduction in metabolic rate compared to normoxia. ArterialPo2was maintained throughout flights. Mixed venous PO2 decreased during the initial portion of flights in hypoxia, indicative of increased tissue O2 extraction. We also discovered that mixed venous temperature decreased during flight, which may significantly increase oxygen loading to hemoglobin.

The distributions of migratory species in the ocean span local, national and international jurisdictions. Across these ecologically interconnected regions, migratory marine species interact with anthropogenic stressors throughout their lives. Migratory connectivity, the geographical linking of individuals and populations throughout their migratory cycles, influences how spatial and temporal dynamics of stressors affect migratory animals and scale up to influence population abundance, distribution and species persistence. Population declines of many migratory marine species have led to calls for connectivity knowledge, especially insights from animal tracking studies, to be more systematically and synthetically incorporated into decision-making. Inclusion of migratory connectivity in the design of conservation and management measures is critical to ensure they are appropriate for the level of risk associated with various degrees of connectivity. Three mechanisms exist to incorporate migratory connectivity into international marine policy which guides conservation implementation: site-selection criteria, network design criteria and policy recommendations. Here, we review the concept of migratory connectivity and its use in international policy, and describe the Migratory Connectivity in the Ocean system, a migratory connectivity evidence-base for the ocean. We propose that without such collaboration focused on migratory connectivity, efforts to effectively conserve these critical species across jurisdictions will have limited effect.

Rescue, rehabilitation and reintroduction programmes are a vital conservation tool used to protect individuals under immediate threat, and then re-establish viable populations in protected habitats. The critically endangered orangutan (Pongo spp.) has experienced significant population decline, primarily due to deforestation and hunting, therefore a considerable number of individuals are rescued from fragmented forests or the illegal pet trade. Whilst the protection of the remaining wild populations remains paramount, the substantial number of individuals currently residing in rescue centres means reintroduction is a crucial strategy for reducing the threat of extinction.
Previously, many reintroductions have been ineffective, primarily attributed to inadequate project management, the introduction of unregulated tourism, or due to the individuals’ failure to acquire vital survival skills. Furthermore, the lack of post-release monitoring often inhibits analyses of the post-release progress, survival rates, and the long-term effects of human-facilitated rehabilitation. Therefore, the reintroduction of Bornean orangutans (Pongo pygmaeus wurmbii) into two protected forests, Bukit Baka Bukit Raya National Park and Bukit Batikap Conservation Forest, Central Kalimantan, provided vital opportunities for understanding how early life experiences and rehabilitation shapes post-release behaviour, ultimately aiding the improvement of reintroduction protocols.
Mixed model analyses of data from consistent post-release monitoring revealed that across the first three months post-release, orangutans displayed behavioural irregularities when compared to their fully-wild counterparts. Temporarily, significantly elevated levels of resting and travelling counteracted reduced levels of feeding, attributed to recovery from transportation and stress from reintroduction into a novel habitat. After approximately three months post-release, behaviour stabilised and closely resembled that of wild orangutans, revealing the robust behavioural flexibility of the species.
Evidence is also provided for distinct dispersal strategies following reintroduction, most notably flanged males were more likely to disperse out of the study site, whilst females often displayed site fidelity and established their home range surrounding their release location. Reintroduced individuals often displayed larger home ranges, when compared to wild individuals, likely in response to substantial overlaps between individuals around release locations, and subsequent resource competition.
Overall, reintroductions were deemed successful, however, differences in dietary composition were discovered, most notably, lower fruit consumption, and a subsequent increased reliance on fall-back foods. Dietary differences are likely attributed to insufficient foraging skills, therefore improvements to rehabilitation protocols are provided, vital for increasing post-release success. Ultimately, the success of this reintroduction programme, and the establishment of new populations reinforces the value of reintroduction as an effective tool for conservation.

Global climate change is expected to have major impacts on biodiversity. Sea turtles have temperature-dependent sex determination, and many populations produce highly femalebiased offspring sex ratios, a skew likely to increase further with global warming. We estimated the primary sex ratio at one of the world's largest green turtle Chelonia mydas rookeries in Guinea-Bissau, West Africa, and explored its resilience to climate change. In 2013 and 2014, we deployed data loggers recording nest (n = 101) and sand (n = 30) temperatures, and identified hatchling sex by histological examination of gonads. A logistic curve was fitted to the data to allow predictions of sex ratio across habitats and through the nesting season. The population-specific pivotal temperature was 29.4°C, with both sexes produced within incubation temperatures from 27.6 to 31.4°C: the transitional range of temperatures (TRT). Primary sex ratio changed from maleto female-biased across relatively small temporal and spatial scales. Overall it was marginally female-biased, but we estimated an exceptionally high male hatchling production of 47.7% (95% CI: 36.7-58.3%) and 44.5% (95% CI: 33.8-55.4%) in 2013 and 2014, respectively. Both the temporal and spatial variation in incubation conditions and the wide range of the TRT suggest resilience and potential for adaptation to climate change if the present nesting habitat remains unchanged. These findings underline the importance of assessing site-specific parameters to understand populations' responses to climate change, particularly with regard to identifying rookeries with high male hatchling production that may be key for the future conservation of sea turtles under projected global warming scenarios.

Birds that migrate across high altitude mountain ranges are faced with the challenge of maintaining vigorous exercise in environments with limited oxygen. Ruddy shelducks are known to use wintering grounds south of the Tibetan Plateau at sea level and breeding grounds north of Himalayan mountain range. Therefore, it is likely these shelducks are preforming high altitude migrations. In this study we analyse satellite telemetry data collected from 15 ruddy shelduck from two populations wintering south of the Tibetan Plateau from 2007 to 2011. During north and south migrations ruddy shelduck travelled 1481 km (range 548–2671 km) and 1238 km (range 548–2689 km) respectively. We find mean maximum altitudes of birds in flight reached 5590 m (range of means 4755–6800 m) and mean maximum climb rates of 0.45 m s–1 (range 0.23–0.74 m s–1). The ruddy shelduck is therefore an extreme high altitude migrant that has likely evolved a range of physiological adaptations in order to complete their migrations.

© 2016 John Wiley & Sons Ltd. Aims: Effective conservation of threatened or endangered species requires a robust understanding of their spatio-temporal distribution. Although a huge amount is known about the movements of Atlantic adult sea turtles, much less is known about juvenile turtles, and much of the life history model is therefore inferred. We set out to describe the spatio-temporal distribution of juvenile loggerheads turtles found around the Canary Islands. Location: Eastern North Atlantic Ocean. Methods: Between 1999 and 2012, we satellite-tracked 24 healthy large juvenile loggerhead turtles (mean straight carapace length = 47.4 cm, range = 34.6-54.5 cm) captured in the waters around the Canary Islands. We describe their regional distribution, identify high-use areas and create a model for habitat suitability using minimum convex polygons, density rasters and ensemble ecological niche modelling, integrated with physical and biological environmental data. Results: Turtles used a huge oceanic area (2.5 million km2) with particularly high usage around the Canary Islands, Spain, Portugal, Morocco and Western Sahara. In spring and summer, turtles generally moved further north towards the Iberian Peninsula. Ecological niche modelling identified sea surface temperature as the most important contributory variable to the habitat models. We also recorded three juvenile turtles making westward migrations away from the eastern Atlantic Ocean, presumably back towards their original natal beaches near sexual maturity. Main conclusions: the results of the present study provide insight into a significant knowledge gap on the spatio-temporal distribution of large juvenile loggerhead turtles in the eastern Atlantic Ocean. The data highlight that turtles occupy a vast open oceanic area, which hampers the ability of static conservation approaches to afford effective protection. However, ensemble ecological niche modelling highlights key suitable habitat for juvenile loggerhead turtles, which could be used in dynamic conservation protection.

Understanding spatial and temporal habitat-use patterns to protect both foraging and breeding grounds of species of concern is crucial for successful conservation. Saona Island in Del Este National Park (DENP), south-eastern Dominican Republic (DR), hosts the only major hawksbill (Eretmochelys imbricata) nesting area in the DR (100 nests yr-1, SD = 8.4, range = 93-111), with the population having been critically reduced through hunting. We satellite tracked 9 female hawksbill turtles, and present analyses of their core-use areas with respect to Marine Protected Areas (MPAs) in both their internesting and foraging areas. Kernel utilization distributions indicated that during the internesting period all turtles remained close to their nesting beaches in small home ranges in the territorial waters of the DR, mostly over the continental shelf (

The integration of satellite telemetry, remotely sensed environmental data, and habitat/environmental modelling has provided for a growing understanding of spatial and temporal ecology of species of conservation concern. The Republic of Cape Verde comprises the only substantial rookery for the loggerhead turtle Caretta caretta in the eastern Atlantic. A size related dichotomy in adult foraging patterns has previously been revealed for adult sea turtles from this population with a proportion of adults foraging neritically, whilst the majority forage oceanically. Here we describe observed habitat use and employ ecological niche modelling to identify suitable foraging habitats for animals utilising these two distinct behavioural strategies. We also investigate how these predicted habitat niches may alter under the influence of climate change induced oceanic temperature rises. We further contextualise our niche models with fisheries catch data and knowledge of fisheries 'hotspots' to infer threat from fisheries interaction to this population, for animals employing both strategies. Our analysis revealed repeated use of coincident oceanic habitat, over multiple seasons, by all smaller loggerhead turtles, whilst larger neritic foraging turtles occupied continental shelf waters. Modelled habitat niches were spatially distinct, and under the influence of predicted sea surface temperature rises, there was further spatial divergence of suitable habitats. Analysis of fisheries catch data highlighted that the observed and modelled habitats for oceanic and neritic loggerhead turtles could extensively interact with intensive fisheries activity within oceanic and continental shelf waters of northwest Africa. We suggest that the development and enforcement of sustainable management strategies, specifically multi-national fisheries policy, may begin to address some of these issues; however, these must be flexible and adaptive to accommodate potential range shift for this species.

Oceanographic fronts are physical interfaces between water masses that differ in properties such as temperature, salinity, turbidity and chlorophyll a enrichment. Bio-physical coupling along fronts can lead to the development of pelagic biodiversity hotspots. A diverse range of marine vertebrates have been shown to associate with fronts, using them as foraging and migration habitats. Elucidation of the ecological significance of fronts generates a better understanding of marine ecosystem functioning, conferring opportunities to improve management of anthropogenic activities in the oceans. This study presents novel insights into the oceanographic drivers of habitat use in a population of marine turtles characterised by an oceanic-neritic foraging dichotomy. Using satellite tracking data from adult female loggerhead turtles Caretta caretta nesting at Cape Verde (n = 12), we tested the hypothesis that oceanic-foraging loggerheads associate with mesocale (10s to 100s of km) thermal fronts. We used high-resolution (1 km) composite front mapping to characterise frontal activity in the Canary Current Large Marine Ecosystem over 2 temporal scales: (1) seasonal front frequency and (2) 7 d front metrics. Our use-availability analysis indicated that oceanic loggerheads show a preference for the highly productive upwelling region between Cape Verde and mainland Africa, an area of intense frontal activity. Within the upwelling region, turtles appear to forage epipelagically around mesoscale thermal fronts, exploiting profitable foraging opportunities resulting from physical aggregation of prey.

Population connectivity is an important consideration in studies of disease transmission and biological conservation, especially with regard to migratory species. Determining how and when different subpopulations intermingle during different phases of the annual cycle can help identify important geographical regions or features as targets for conservation efforts and can help inform our understanding of continental-scale disease transmission. In this study, stable isotopes of hydrogen and carbon in contour feathers were used to assess the degree of molt-site fidelity among Bar-headed Geese (Anser indicus) captured in north-central Mongolia. Samples were collected from actively molting Bar-headed Geese (n = 61), and some individual samples included both a newly grown feather (still in sheath) and an old, worn feather from the bird's previous molt (n = 21). Although there was no difference in mean hydrogen isotope ratios for the old and new feathers, the isotopic variance in old feathers was approximately three times higher than that of the new feathers, which suggests that these birds use different and geographically distant molting locations from year to year. To further test this conclusion, online data and modeling tools from the isoMAP website were used to generate probability landscapes for the origin of each feather. Likely molting locations were much more widespread for old feathers than for new feathers, which supports the prospect of low molt-site fidelity. This finding indicates that population connectivity would be greater than expected based on data from a single annual cycle, and that disease spread can be rapid even in areas like Mongolia where Bar-headed Geese generally breed in small isolated groups.

Geese need to hug the land to fly high
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. Animal migrations provide numerous examples of astonishing feats. Impressive even among these is the migration of bar-headed geese across the Himalayan Mountains, which reach heights of thousands of meters. Bishop
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. remotely monitored birds' heart rates, movement, and body temperature during migration. The geese “hug” the landforms, taking advantage of drafting and wind patterns. This unexpected strategy conserves energy, even though it means the geese repeatedly lose, and must then regain, altitude.
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Management of species of conservation concern requires knowledge of demographic parameters, such as rates of recruitment, survival, and growth. In the Caribbean, hawksbill turtles (Eretmochelys imbricata) have been historically exploited in huge numbers to satisfy trade in their shells and meat. In the present study, we estimated growth rate of juvenile hawksbill turtles around Anegada, British Virgin Islands, using capture-mark-recapture of 59 turtles over periods of up to 649 days. Turtles were recaptured up to six times, having moved up to 5.9 km from the release location. Across all sizes, turtles grew at an average rate of 9.3 cm year-1 (range 2.3-20.3 cm year-1), and gained mass at an average of 3.9 kg year-1 (range 850 g-16.1 kg year-1). Carapace length was a significant predictor of growth rate and mass gain, but there was no relationship between either variable and sea surface temperature. These are among the fastest rates of growth reported for this species, with seven turtles growing at a rate that would increase their body size by more than half per year (51-69% increase in body length). This study also demonstrates the importance of shallow water reef systems for the developmental habitat for juvenile hawksbill turtles. Although growth rates for posthatching turtles in the pelagic, and turtles larger than 61 cm, are not known for this population, the implications of this study are that Caribbean hawksbill turtles in some areas may reach body sizes suggesting sexual maturity in less time than previously considered. In the present study, we estimated growth rate of juvenile hawksbill turtles around Anegada, British Virgin Islands, using capture-mark-recapture of 59 turtles over periods of up to 649 days. Across all sizes, turtles grew at an average rate of 9.3 cm per year (range 2.3-20.3 cm year-1), and gained mass at an average of 3.9 kg year-1 (range 850 g-16.1 kg year-1). These are among the fastest rates of growth reported for this species, with seven turtles growing at a rate that would increase their body size by more than half per year (51-69% increase in body length). © 2014 the Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

Summary: 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. 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. 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. 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. 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. 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. 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.

Nesting by three species of marine turtles persists in the Dominican Republic, despite historic threats and long-term population decline. We conducted a genetic survey of marine turtles in the Dominican Republic in order to link them with other rookeries around the Caribbean. We sequenced a 740bp fragment of the control region of the mitochondrial DNA of 92 samples from three marine turtle species [hawksbill (n = 48), green (n = 2) and leatherback (n = 42)], and incorporated published data from other nesting populations and foraging grounds. The leatherback turtle (Dermochelys coriacea) in the Dominican Republic appeared to be isolated from Awala-Yalimapo, Cayenne, Trinidad and St. Croix but connected with other Caribbean populations. Two distinct nesting populations of hawksbill turtles (Eremochelys imbricata) were detected in the Dominican Republic and exhibited interesting patterns of connectivity with other nesting sites and juvenile and adult male foraging aggregations. The green sea turtle (Chelonia mydas) has almost been extirpated from the Dominican Republic and limited inference could be made from our samples. Finally, results were compared with Lagrangian drifting buoys and published Lagrangian virtual particles that travelled through the Dominican Republic and Caribbean waters. Conservation implications of sink-source effects or genetic isolation derived from these complex inter-connections are discussed for each species and population. © 2013 Carreras et al.

Evolutionary theory predicts that male and female offspring should be produced at a 1:1 ratio, but this may rarely be the case for species in which sex is determined during incubation by temperature, such as marine turtles. Estimates of primary sex ratio suggest that marine turtle sex ratios are highly skewed, with up to 9 females per male. We captured juvenile hawksbill turtles Eretmochelys imbricata in waters around Anegada, British Virgin Islands, a regionally important foraging aggregation, and analysed concentrations of plasma testosterone and oestradiol- 17β from 62 turtles to estimate sex ratio. There were 2.4 to 7.7 times more females than males. Testosterone concentrations correlated with sampling date and sea surface temperature (SST), with higher con centrations in the late summer when SST was highest, suggesting that assigning sex through threshold values of sex hormones must be carried out cautiously. The sex ratio in the juvenile foraging aggregation around Anegada is more male biased than at other locations, suggesting that turtles at Anegada have resilience against feminising effects of climate change. Future work should (1) integrate the relative contributions of different genetic stocks to foraging aggregations and (2) investigate the annual and seasonal cycles of sex hormones, and differences among individuals and life history stages. © Inter-Research 2013.

From 1998 to 2008, 68 adult female loggerhead sea turtles (Caretta caretta) were instrumented with platform transmitter terminals at nesting beaches in Georgia, North Carolina (NC) and South Carolina (SC) on the East Coast of the United States of America (30°48′N, 81°28′W to 33°51′N, 77°59′W). The majority of post-nesting loggerheads (N = 42, 62 %) migrated to foraging habitats in the Mid-Atlantic Bight during May-October, with a subsequent migration occurring during November-March to foraging habitats south of Cape Hatteras, NC. Nine (13 %) loggerheads initially foraged in the near-shore, coastal areas of the South Atlantic Bight, but moved to offshore habitats-closer to the Gulf Stream-during November-March, while fourteen (21 %) loggerheads remained in foraging areas along the mid-continental shelf off of the eastern coast of Florida and/or continued southward to Florida Bay and the Bahamas. The present study delineates important, post-nesting foraging habitats and migration corridors where loggerheads may interact with commercial fisheries-providing managers opportunities to develop and implement optimally effective conservation actions for the recovery of this threatened species. © 2013 Springer-Verlag Berlin Heidelberg.

In recent years, information about the movements and timing of migration by male sea turtles has begun to be unraveled. Here, we present the first satellite tracking of male loggerhead sea turtles (Caretta caretta) in the eastern Atlantic. Satellite linked transmitters were attached to five adult males, captured in the near shore waters off Boavista, Republic of Cape Verde. This archipelago hosts the single most important breeding site of loggerhead turtles in the eastern Atlantic. Animals were tracked for periods ranging between 48 and 537. days, including a probable annual remigration to the vicinity of the nesting ground for one turtle. Males showed a variety of movement patterns both during and after the breeding season. of three males that transmitted for 85, 329 and 537. days, two (the smallest) migrated east and remained in oceanic waters for the tracking period and another (larger turtle) migrated 810. km northeast, to neritic waters off the coast of Mauritania, Western Africa. Results suggest males may show the same size-linked dichotomy in migratory strategies, as has been shown for females from this population. © 2013 Elsevier B.V.

Bar-headed geese are renowned for migratory flights at extremely high altitudes over the world's tallest mountains, the Himalayas, where partial pressure of oxygen is dramatically reduced while flight costs, in terms of rate of oxygen consumption, are greatly increased. Such a mismatch is paradoxical, and it is not clear why geese might fly higher than is absolutely necessary. In addition, direct empirical measurements of high-altitude flight are lacking. We test whether migrating bar-headed geese actually minimize flight altitude and make use of favourable winds to reduce flight costs. By tracking 91 geese, we show that these birds typically travel through the valleys of the Himalayas and not over the summits. We report maximum flight altitudes of 7290 m and 6540 m for southbound and northbound geese, respectively, but with 95 per cent of locations received from less than 5489 m. Geese travelled along a route that was 112 km longer than the great circle (shortest distance) route, with transit ground speeds suggesting that they rarely profited from tailwinds. Bar-headed geese from these eastern populations generally travel only as high as the terrain beneath them dictates and rarely in profitable winds. Nevertheless, their migration represents an enormous challenge in conditions where humans and other mammals are only able to operate at levels well below their sea-level maxima.

It is now well understood that climate change has the potential to dramatically affect biodiversity, with effects on spatio-temporal distribution patterns, trophic relationships and survivorship. In the marine turtles, sex is determined by incubation temperature, such that warming temperatures could lead to a higher production of female hatchlings. By measuring nest temperature, and using a model to relate the incubation temperature to sex ratio, we estimate that Caribbean Colombian leatherback sea turtles currently produce approximately 92% female hatchlings. We modelled the relationship between incubation, sand and air temperature, and under all future climate change scenarios (0.4-6.0 °C warming over the next 100 years), complete feminization could occur, as soon as the next decade. However, male producing refugia exist in the periphery of smaller nests (0.7 °C cooler at the bottom than at the centre), within beaches (0.3 °C cooler in the vegetation line and inter-tidal zone) and between beaches (0.4 °C higher on dark beaches), and these natural refugia could be assigned preferential conservation status. However, there exists a need to develop strategies that may ameliorate deleterious effects of climate-induced temperature changes in the future. We experimentally shaded clutches using screening material, and found that it was effective in reducing nest temperature, producing a higher proportion of male hatchlings, without compromising the fitness or hatching success. Artificial shade in hatcheries is a very useful and simple tool in years or periods of high environmental temperatures. Nevertheless, this is only an emergency response to the severe impacts that will eventually have to be reversed if we are to guarantee the stability of the populations. © 2011 Blackwell Publishing Ltd.

The advent of telemetry has improved knowledge of the spatio-temporal distribution of marine species of conservation concern. Among the sea turtles, the movements of the hawksbill turtle Eretmochelys imbricata are among the least well described. We tracked 10 adult female hawksbill turtles by satellite after nesting in the Dominican Republic (DR) and describe a dichotomy in patterns of movement: some (n = 2) turtles remained in the DR, while others migrated to waters off Honduras and Nicaragua (n = 5) and the Bahamas (n = 1). Transmitters on 2 turtles failed during migration, before they reached their final foraging grounds. We present results from long tracking durations for 3 turtles, including 3 entire remigration intervals, high-lighting foraging ground and nest-site fidelity. Threats to hawksbill turtles are not well documented for Nicaragua or neighbouring Honduras and represent a major information gap. We suggest that directing conservation efforts to regionally important foraging areas, such as those in Nicaragua, and strengthening national conservation in each nation with significant hawksbill nesting offers a clear way forward for the conservation of hawksbill turtles in the region. © Inter-Research 2012 · www.int-res.com.

Conserving and managing populations of marine vertebrates can be complex when they occupy the waters of multiple nations, crossing heterogeneous legal and management landscapes. Hawksbill turtles (. Eretmochelys imbricata) are subject to varying levels of use in Caribbean countries and their conservation and management is complicated by the extent to which they are a 'shared' resource. In 1997 and 2000, Cuba attempted to 'downlist' hawksbills from Cuban waters to CITES Appendix II to allow limited international trade. The research on movement and dispersal of hawksbills reported here was undertaken to better inform discussion about the impacts of their harvest. Flipper tagging and satellite tracking demonstrate that the majority of study turtles remained in Cuban territorial waters. of 1170 hawksbills tagged (525 adults and 606 juveniles), 12% (. n=. 143) were recaptured. All recaptured adults (. n=. 16 males, 38 nesting females and 30 adult females in-water) were in Cuban waters. of the 59 juveniles recaptured, only four recaptures were outside Cuban waters (Nicaragua. =. 2, Colombia. =. 1, USA. =. 1). Fourteen hawksbills tagged in the waters of other nations were recaptured in Cuban waters. We also satellite tracked 21 turtles (one adult male, ten nesting females and ten non-nesting adult females), of which five tags failed, 11 stayed in Cuban waters for the duration of transmissions (1-809. days) and five foraged in the waters of other nations (Mexico. n=. 1 an adult female; Honduras. n=. 2, both post-nesting turtles; Colombia. n=. 1 an adult female; and the eastern Lesser Antilles. n=. 1, an adult male), with differences for nesting and non-nesting turtles. Our results, demonstrating extended site fidelity within Cuban waters, suggest that strengthening national management within national jurisdictions that host hawksbill turtles is fundamental to improving regional conservation as a whole. © 2012 Elsevier Ltd.

We describe the real-time movements of the last of the marine mega-vertebrate taxa to be satellite tracked - the giant manta ray (or devil fish, Manta birostris), the world's largest ray at over 6 m disc width. Almost nothing is known about manta ray movements and their environmental preferences, making them one of the least understood of the marine mega-vertebrates. Red listed by the International Union for the Conservation of Nature as 'Vulnerable' to extinction, manta rays are known to be subject to direct and incidental capture and some populations are declining. Satellite-tracked manta rays associated with seasonal upwelling events and thermal fronts off the Yucatan peninsula, Mexico, and made short-range shuttling movements, foraging along and between them. The majority of locations were received from waters shallower than 50 m deep, representing thermally dynamic and productive waters. Manta rays remained in the Mexican Exclusive Economic Zone for the duration of tracking but only 12% of tracking locations were received from within Marine Protected Areas (MPAs). Our results on the spatio-temporal distribution of these enigmatic rays highlight opportunities and challenges to management efforts.

Birds that fly over mountain barriers must be capable of meeting the increased energetic cost of climbing in low-density air, even though less oxygen may be available to support their metabolism. This challenge is magnified by the reduction in maximum sustained climbing rates in large birds. Bar-headed geese (Anser indicus) make one of the highest and most iconic transmountain migrations in the world. We show that those populations of geese that winter at sea level in India are capable of passing over the Himalayas in 1 d, typically climbing between 4,000 and 6,000 m in 7-8 h. Surprisingly, these birds do not rely on the assistance of upslope tailwinds that usually occur during the day and can support minimum climb rates of 0.8-2.2 km·h(-1), even in the relative stillness of the night. They appear to strategically avoid higher speed winds during the afternoon, thus maximizing safety and control during flight. It would seem, therefore, that bar-headed geese are capable of sustained climbing flight over the passes of the Himalaya under their own aerobic power.

Understanding the spatial movements of threatened marine species, such as sea turtles, is essential as a means of informing appropriate conservation management. Although novel techniques for tracking spatial movements are becoming more widely available (such as satellite tracking), simple techniques such as mark-release-recapture remain effective. A flipper tagging and recovery program in Cuba tagged 210 loggerhead turtles over 14 yr and recovered 7% of the tags between 2 d and 3 yr later (mean = 296 d). All but one turtle was recaptured in Cuban waters, and data showed limited movement of turtles between northern and southern coasts. A further 50 turtles were recovered that had been tagged in foreign projects, the majority of which were from the USA (but also Mexico, the Bahamas, Canary Islands and Spain). A range of life stages of loggerhead turtles are found in Cuban waters year-round, and given that Cuba has the second largest reef in the Caribbean, it likely provides foraging habitat for significant numbers of loggerhead turtles from at least 6 different countries. © Inter-Research 2010.

Marine turtles utilise terrestrial and marine habitats and several aspects of their life history are tied to environmental features that are altering due to rapid climate change. We overview the likely impacts of climate change on the biology of these species, which are likely centred upon the thermal ecology of this taxonomic group. Then, focusing in detail on three decades of research on the loggerhead turtle (Caretta caretta L.), we describe how much progress has been made to date and how future experimental and ecological focus should be directed. Key questions include: what are the current hatchling sex ratios from which to measure future climate-induced changes? What are wild adult sex ratios and how many males are necessary to maintain a fertile and productive population? How will climate change affect turtles in terms of their distribution?

Virulent outbreaks of highly pathogenic avian influenza (HPAI) since 2005 have raised the question about the roles of migratory and wild birds in the transmission of HPAI. Despite increased monitoring, the role of wild waterfowl as the primary source of the highly pathogenic H5N1 has not been clearly established. The impact of outbreaks of HPAI among species of wild birds which are already endangered can nevertheless have devastating consequences for the local and non-local ecology where migratory species are established. Understanding the entangled dynamics of migration and the disease dynamics will be key to prevention and control measures for humans, migratory birds and poultry. Here, we present a spatial dynamic model of seasonal migration derived from first principles and linking the local dynamics during migratory stopovers to the larger scale migratory routes. We discuss the effect of repeated epizootic at specific migratory stopovers for bar-headed geese (Anser indicus). We find that repeated deadly outbreaks of H5N1 on stopovers during the autumn migration of bar-headed geese could lead to a larger reduction in the size of the equilibrium bird population compared with that obtained after repeated outbreaks during the spring migration. However, the opposite is true during the first few years of transition to such an equilibrium. The age-maturation process of juvenile birds which aremore susceptible to H5N1 reinforces this result. © 2010 the Royal Society.

Marine turtles occupy a wide range of terrestrial and marine habitats, and many aspects of their life history have been demonstrated to be closely tied to climatic variables such as ambient temperature and storminess. As a group, therefore, marine turtles may be good indicators of climate change effects on coastal and marine habitats. Despite the small number of species in the taxon and a growing body of research in the field, the evidence base to predict resultant impacts of climate change remains relatively poor. We review the data from peer-reviewed publications to assess the likely impacts of climate change on marine turtles and highlight the types of data that would be most useful for an accurate assessment of future effects. The cumulative indications from these previous studies indicate that future research should focus on: (1) climate change effects on key habitats upon which turtles depend; (2) factors that influence nest site selection; (3) the consequences of skewed primary sex ratios; and (4) the effect of climate change on turtles at sea, for example range shifts and dietary breadth. Although it is too early to give detailed management recommendations, careful protection of coastlines along which turtles nest should be considered, as should the protection of beaches that produce male hatchlings, which may be of increased importance in the future. More active management approaches, for example translocation of eggs to suitable yet vacant nesting beaches, may be necessary to consider under worst-case scenarios. © Inter-Research 2009.

The connectivity and frequency of exchange between sub-populations of migratory birds is integral to understanding population dynamics over the entire species' range. True geese are highly philopatric and acquire lifetime mates during the winter, suggesting that the number of distinct sub-populations may be related to the number of distinct wintering areas. In the Bar-headed Goose Anser indicus, a species found exclusively in Central Asia, the connectivity between breeding and wintering areas is not well known. Their migration includes crossing a broad front of the Himalaya Cordillera, a significant barrier to migration for most birds. Many Bar-headed Geese fly to breeding areas on the Tibetan-Qinghai Plateau (TQP), the highest plateau in the world. From 2005-2008, 60 Bar-headed Geese were captured and marked with satellite transmitters in Nepal (n = 2), India (n = 6), China (n = 29), and Mongolia (n = 23) to examine their migration and distribution. Distinct differences were observed in their migration corridors and timing of movements, including an apparent leap-frog migration pattern for geese from Mongolia. Measurements of geese from Mongolia were larger than their counterparts from China, providing some evidence of morphological differences. Alteration of habitats in China, including the warming effects of climate change on glaciers increasing runoff to TQP wetlands, may be changing goose migration patterns and timing. With the exception of one individual, all geese from Qinghai Lake, China wintered in the southern TQP near Lhasa, and their increasing numbers in that region may be related to the effects of climate change and agricultural development. Thus, our findings document both morphological and geographical variation in sub-populations of Bar-headed Geese, but their resilience to environmental change may be lost if migratory short-stopping results in larger congregations restricted to a smaller number of wintering areas. © Wildfowl & Wetlands Trust.