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As climate-related impacts threaten marine biodiversity globally, it is important to adjust conservation efforts to mitigate the effects of climate change. Translating scientific knowledge into practical management, however, is often complicated due to resource, economic and policy constraints, generating a knowledge-action gap. To develop potential solutions for marine turtle conservation, we explored the perceptions of key actors across 18 countries in the Mediterranean. These actors evaluated their perceived relative importance of 19 adaptation and mitigation measures that could safeguard marine turtles from climate change. Of importance, despite differences in expertise, experience and focal country, the perceptions of researchers and management practitioners largely converged with respect to prioritizing adaptation and mitigation measures. Climate change was considered to have the greatest impacts on offspring sex ratios and suitable nesting sites. The most viable adaptation/mitigation measures were considered to be reducing other pressures that act in parallel to climate change. Ecological effectiveness represented a key determinant for implementing proposed measures, followed by practical applicability, financial cost, and societal cost. This convergence in opinions across actors likely reflects long-standing initiatives in the Mediterranean region towards supporting knowledge exchange in marine turtle conservation. Our results provide important guidance on how to prioritize measures that incorporate climate change in decision-making processes related to the current and future management and protection of marine turtles at the ocean-basin scale, and could be used to guide decisions in other regions globally. Importantly, this study demonstrates a successful example of how interactive processes can be used to fill the knowledge-action gap between research and management., This work was conducted under FutureMares EU project that received funding from the European Union's Horizon 2020 research and innovation programme under grant agreement No 869300. The Mediterranean Marine Turtle Working Group was established in 2017 and is continuously supported by MedPAN and the National Marine Park of Zakynthos. The work of AC was supported by the Hellenic Foundation for Research and Innovation (H.F.R.I.) under the “First Call for H.F.R.I. Research Projects to support Faculty members and Researchers and the procurement of high-cost research equipment grant” (Project Number: 2340).

Aim: Understanding the spatial ecology of animal movements is a critical element in conserving long-lived, highly mobile marine species. Analyzing networks developed from movements of six sea turtle species reveals marine connectivity and can help prioritize conservation efforts. Location: Global. Methods: We collated telemetry data from 1235 individuals and reviewed the literature to determine our dataset's representativeness. We used the telemetry data to develop spatial networks at different scales to examine areas, connections, and their geographic arrangement. We used graph theory metrics to compare networks across regions and species and to identify the role of important areas and connections. Results: Relevant literature and citations for data used in this study had very little overlap. Network analysis showed that sampling effort influenced network structure, and the arrangement of areas and connections for most networks was complex. However, important areas and connections identified by graph theory metrics can be different than areas of high data density. For the global network, marine regions in the Mediterranean had high closeness, while links with high betweenness among marine regions in the South Atlantic were critical for maintaining connectivity. Comparisons among species-specific networks showed that functional connectivity was related to movement ecology, resulting in networks composed of different areas and links. Main conclusions: Network analysis identified the structure and functional connectivity of the sea turtles in our sample at multiple scales. These network characteristics could help guide the coordination of management strategies for wide-ranging animals throughout their geographic extent. Most networks had complex structures that can contribute to greater robustness but may be more difficult to manage changes when compared to simpler forms. Area-based conservation measures would benefit sea turtle populations when directed toward areas with high closeness dominating network function. Promoting seascape connectivity of links with high betweenness would decrease network vulnerability. © 2022 The Authors. Diversity and Distributions published by John Wiley & Sons Ltd.

Machine learning algorithms are being increasingly used to process large volumes of wildlife imagery data from unmanned aerial vehicles (UAVs); however, suitable algorithms to monitor multiple species are required to enhance efficiency. Here, we developed a machine learning algorithm using a low‐cost computer. We trained a convolutional neural network and tested its performance in: (1) distinguishing focal organisms of three marine taxa (Australian fur seals, loggerhead sea turtles and Australasian gannets; body size ranges: 0.8–2.5 m, 0.6–1.0 m, and 0.8–0.9 m, respectively); and (2) simultaneously delineating the fine‐scale movement trajectories of multiple sea turtles at a fish cleaning station. For all species, the algorithm performed best at detecting individuals of similar body length, displaying consistent behaviour or occupying uniform habitat (proportion of individuals detected, or recall of 0.94, 0.79 and 0.75 for gannets, seals and turtles, respectively). For gannets, performance was impacted by spacing (huddling pairs with offspring) and behaviour (resting vs. flying shapes, overall precision: 0.74). For seals, accuracy was impacted by morphology (sexual dimorphism and pups), spacing (huddling and creches) and habitat complexity (seal sized boulders) (overall precision: 0.27). For sea turtles, performance was impacted by habitat complexity, position in water column, spacing, behaviour (interacting individuals) and turbidity (overall precision: 0.24); body size variation had no impact. For sea turtle trajectories, locations were estimated with a relative positioning error of

In a warming climate, male sea turtles may become increasingly rare due to temperature-dependent sex determination with females being produced at warmer temperatures. Hence there is widespread concern that a lack of adult males may impact population viability. However, there is controversy over this scenario and here we review aspects of the biology of male sea turtles that will help mitigate female-biased hatchling sex ratios. In particular, there is strong evidence that males generally breed more frequently than females (i.e. have a shorter remigration interval) and that individual breeding males actively search for females and may mate with multiple females from different nesting sites. These aspects of the biology of male turtles will cause female-biased hatchling sex ratios to translate into more balanced adult sex ratios on the breeding grounds (i.e. operational sex ratios). Sexual dimorphism is widespread with adult male turtles generally being smaller than females. In freshwater turtles, this sexual dimorphism is linked to earlier age at maturity for males, although this possibility has not been examined widely in sea turtles. We make a forward-looking horizon-scanning prediction for key changes that might be expected at sea turtle breeding grounds if female-biased sex ratios become so extreme that male turtles start to become limiting and start driving populations to extinction. In particular, as the numbers of adult males on the breeding grounds become limiting there may be changes in egg fertility, multiple paternity and hatching success within clutches.

Documenting how human pressure on wildlife changes over time is important to minimise potential adverse effects through implementing appropriate management and policy actions; however, obtaining objective measures of these changes and their potential impacts is often logistically challenging, particularly in the natural environment. Here, we developed a modular stochastic model that infers the ratio of actual viewing pressure on wildlife in consecutive time periods (years) using social media, as this medium is widespread and easily accessible. Pressure was calculated from the number of times individual animals appeared in social media in pre-defined time windows, accounting for time-dependent variables that influence them (e.g. number of people with access to social media). Formulas for the confidence intervals of viewing pressure ratios were rigorously developed and validated, and corresponding uncertainty was quantified. We applied the developed framework to calculate changes to wildlife viewing pressure on loggerhead sea turtles (Caretta caretta) at Zakynthos island (Greece) before and during the COVID-19 pandemic (2019-2021) based on 2646 social media entries. Our model ensured temporal comparability across years of social media data grouped in time window sizes, by correcting for the interannual increase of social media use. Optimal sizes for these windows were delineated, reducing uncertainty while maintaining high time-scale resolution. The optimal time window was around 7-days during the peak tourist season when more data were available in all three years, and >15 days during the low season. In contrast, raw social media data exhibited clear bias when quantifying changes to viewing pressure, with unknown uncertainty. The framework developed here allows widely-available social media data to be used objectively when quantifying temporal changes to wildlife viewing pressure. Its modularity allowed viewing pressure to be quantified for all data combined, or subsets of data (different groups, situations or locations), and could be applied to any site supporting wildlife exposed to tourism.

There is an intense interest in long-term trends of species abundance that may reflect, for example, climate change or conservation actions. Less well studied are patterns in the magnitude of inter-annual variability in abundance across large spatial scales. We collated abundance time-series for 133 nesting sites across the globe of the seven sea turtle species. Inter-annual variability in nest numbers was lowest in loggerhead turtles (Caretta caretta), Kemp’s ridley (Lepidochelys kempii) and flatback turtle (Natator depressus) and highest in green turtles (Chelonia mydas), likely reflecting their lower trophic position compared to other species and hence tighter coupling of food availability to environmental conditions each year. The annual number of nests in green turtles could vary by 60-fold between successive years. We identified regional patterns in the magnitude of inter-annual variability in green turtle nest numbers, variability being highest for nesting beaches around Australia and lowest in the western Indian Ocean and equatorial Atlantic. These regional patterns are likely linked to corresponding patterns of environmental variability with, for example, areas subjected environmental extremes as part of the El Nino Southern Oscillation (ENSO) showing high inter-annual variability in nest numbers.

Establishing how wildlife viewing pressure is distributed across individual animals within a population can inform the management of this activity, and ensure targeted individuals or groups are sufficiently protected. Here, we used social media data to quantify whether tourism pressure varies in a loggerhead sea turtle (Caretta caretta) population and elucidate potential implications. Laganas Bay (Zakynthos, Greece) supports both breeding (migratory, and hence transient) and foraging (resident) turtles, with turtle viewing representing a major component of the tourism industry. Social media entries spanning two seasons (April to November, 2018 and 2019) were evaluated, and turtles were identified via photo-identification. For both years, 1684 and 2105 entries of 139 and 122 unique turtles were obtained from viewings, respectively (boats and underwater combined). However, while residents represented less than one-third of uniquely identified turtles, they represented 81.9% and 87.9% of all entries. Even when the seasonal breeding population was present (May to July), residents represented more than 60% entries. Of note, the same small number of resident turtles (

Tracking of juvenile sea turtles is a research priority to inform the protection of relevant habitats and ensure sustainable rates of recruitment into adult populations. Based on satellite tracking, mixed stock analysis, and mark–recapture studies, Drini Bay in the South Adriatic Sea, Central Mediterranean, has been confirmed as an important foraging site used by loggerhead turtles from all major rookeries in the Mediterranean subpopulation. Three juvenile loggerhead turtles (Caretta caretta) from Drini Bay were tracked for 763, 364, and 211 days respectively. All turtles exhibited different movement patterns. The two smaller turtles ranged widely beyond the bay. One of these (69.5 cm curved‐carapace length; CCL) used the coldest region of the Mediterranean during the first winter of tracking where mean weekly temperatures dipped below 12°C but moved southwards to warmer waters during the second winter. In comparison, the other (66 cm CCL) individual moved south to winter in warmer waters from the outset. Both individuals returned to Drini Bay during summer, demonstrating fidelity to the study site. The third turtle, which was larger (76 cm CCL), remained in Drini Bay for the duration of tracking. These results support the findings of other mark–recapture studies at Drini Bay, which have suggested that the foraging site is being used in a transient way by immature turtles. All three turtles preferentially used the shallow waters (

Temperature is often used to infer how climate influences wildlife distributions; yet, other parameters also contribute, separately and combined, with effects varying across geographical scales. Here, we used an unoccupied aircraft system to explore how environmental parameters affect the regional distribution of the terrestrial and marine breeding habitats of threatened loggerhead sea turtles (Caretta caretta). Surveys spanned four years and ~620 km coastline of western Greece, encompassing low (

Quantifying the extent to which animals detect and respond to human presence allows us to identify pressure (disturbance) and inform conservation management objectively;however, obtaining baselines against which to compare human impact is hindered in areas where human activities are already well-established. For example, Zakynthos Island (Greece, Mediterranean) receives around 850,000 visitors each summer, while supporting an important loggerhead sea turtle rookery (~300 individuals/season). The coronavirus (COVID-19) driven absence of tourism in May?June 2020 provided an opportunity to evaluate the distribution dynamics of this population in the absence (2020) versus presence (2018 and 2019) of visitors using programmed unmanned aerial system (UAS) surveys. Ambient sea temperature transitioned from suboptimal for breeding in May to optimal in late June, with turtle distribution appearing to shift from shallow (to benefit from waters 3?5°C above ambient) to deeper waters in 2018 and 2019, but not 2020. The 2020 dataset demonstrated that increased tourism pressure, not temperature, drives turtles offshore. Specifically, >50% of turtles remained within 100 m of shore at densities of 25?50 visitors/km, even when sea temperature rose, with 2018 and 2019 data supporting this trend. Reduced access to warmer, nearshore waters by tourism could delay the onset of nesting and increase the length of the egg maturation period between nesting events (internesting interval) at this site. A coastal refuge zone could be delimited in May-June where touristic infrastructure is minimal, but also where turtles frequently aggregate. In conclusion, sea turtles appear capable of perceiving changes in the level of human pressure at fine spatial and temporal scales, and adjusting their distribution accordingly.

There is increasing demand for efficient ways to process large volumes of data from visual-based remote-technology, such as unmanned aerial vehicles (UAVs) in ecology and conservation, with machine learning methods representing a promising avenue to address varying user demands. Here, we evaluated current trends in how machine learning and UAVs are used to process imagery data for detecting animals and vegetation across habitats, placing emphasis on their utility for endangered species. We reviewed 213 publications that used UAVs at 256 study sites, of which just 89 (42%) used machine learning to assess the visual data. We evaluated geographical and temporal trends and identified how each technology is used at a global scale. We also identified the most commonly encountered machine-learning methods, including potential reasons for their limited use in ecology and possible solutions. Thirteen out of the 17 habitats defined by the International Union for Conservation of Nature (IUCN) habitat classification scheme were monitored using UAVs, while 12 habitats were monitored using both UAVs and machine learning. Our results show that, while machine learning is already being used across many habitat types, it is primarily restricted to more uniform habitats at present. Out of 173 plant and animal species monitored using UAV surveys, 30 were of conservation concern, with machine learning being used to assess UAV imagery data for 9 of these species. In conclusion, we anticipate that the joint use of UAVs and machine learning for ecological research and conservation will expand as machine learning methods become more accessible.

International audience; Although transmissible cancers have, so far, only been documented in three independent animal groups, they not only impact animals that have high economic, environmental and social significance, but they are also one of the most virulent parasitic life forms. Currently known transmissible cancers traverse terrestrial and marine environments, and are predicted to be more widely distributed across animal groups; thus, the implementation of effective collaborative scientific networks is important for combating existing and emerging forms. Here, we quantify how collaborative effort on the three known transmissible cancers has advanced through the formation of collaborative networks among institutions and disciplines. These three cancers occur in bivalves (invertebrates—disseminated neoplasia; DN), Tasmanian devils (vertebrate—marsupial; devil facial tumour disease; DFTD) and dogs (vertebrate—eutherian mammal; canine transmissible venereal tumour; CTVT). Research on CTVT and DN has been conducted since 1876 and 1969, respectively, whereas systematic research on DFTD only started in 2006. Yet, collaborative effort on all three diseases is global, encompassing six major Scopus subject areas. Collaborations steadily increased between 1963 and 2006 for CTVT and DN, with similar acceleration for all three cancers since 2006. Network analyses demonstrated that scientists are organizing themselves into efficient collaborative networks; however, these networks appear to be far stronger for DFTD and DN, possibly due to the recent detection of new strains adding impetus to research and associated publications (enhancing citation trajectories). In particular, global and multidisciplinary collaborations formed almost immediately after DFTD research was initiated, leading to similar research effort and relatively greater research outputs compared to the other two diseases. Therefore, in the event of outbreaks of new lineages of existing transmissible cancers, or the discovery of new transmissible cancers in the future, the rapid formation of international collaborations spanning relevant disciplines is vital for the efficient management of these diseases.

Aim Identifying processes that determine climatic niche dynamics is difficult, especially for wide-ranging species where niche conservatism might differ at different life stages. Evolutionary history shapes climatic niche, with populations of the same or phylogenetically close species occupying similar conditions. However, the geographical separation of populations also leads to niche evolution as an adaptive response to local conditions. Here, we use five sea turtle species (loggerhead, green, leatherback, hawksbill, olive ridley) to test whether short- (environmental adaptation) or long-term (phylogenetic separation) evolutionary events determined the level of niche conservatism among distinct groups of nesting populations (termed Regional Management Units; RMUs) distributed across different ocean basins. Location Global. Methods We estimated the climatic niche of 4,829 georeferenced nesting locations using air and sea surface parameters. We quantified niche overlap among RMUs of the same species at a global scale and examined how the geographical ranges of each RMU are correlated with niche similarity. We also investigated the extent of niche conservatism among RMUs of different species in the same ocean basin. Results We found limited within-species niche similarity of RMUs at a global scale, with geographical range having negligible impact (latitudinal overlap, range size asymmetries). We detected generally higher niche overlap between RMUs of different species located within the same ocean basin, where the geographical range had a stronger effect on niche similarity. Main conclusions We show that adaptation to local conditions potentially prevails over distant evolutionary events when considering the climatic niche of RMUs of the same species across different oceans. Because sea turtles, like other long-distance migratory animals, occupy distinct climatic niches, our findings support the importance of understanding this phenomenon for different life stages, populations and species to preserve unique adaptations to different environments that might enhance future viability under climate change, as well as help identify key conservation areas.

Summary Offspring and breeding (operational) sex ratios (OSR) are a key component of demographic studies. While offspring sex ratios are often relatively easy to measure, measuring OSRs is often far more problematic. Yet highly skewed OSRs, and a lack of male-female encounters, may be an important extinction driver. Using loggerhead sea turtles (Caretta caretta) as a case study, we showed the utility of drones, i.e. unmanned aerial vehicles (UAVs) to distinguish adult males and females in a marine breeding area, using a combination of morphological characteristics (tail length) and behavioural differences (active mating, courting and searching by males versus resting by females). Through repeated surveys, we documented seasonal changes in the OSR. While the number, and ratio, of males and females on the breeding grounds changed massively, the ratio of receptive females (derived from the rate of influx of new individuals to the area) to breeding males remained close to 1:1 for much of the period before nesting commenced. Hence, we show how large imbalances in the number of adult males and females may translate into relatively balanced OSRs. Our results suggest that the departure of males from the breeding grounds is linked to a decline in female receptivity, with female sea turtles being known to store sperm to ensure high clutch fertility throughout the nesting season. In conclusion, while we detected up to three times more females than males at the breeding ground, at present, OSRs appear stable. However, because most males breed annually (versus biannually by females), there might only be ˜100 males in the adult population (i.e. adult sex ratio of 1:7.5), which might become further skewed under expected climate change scenarios; thus, we need to identify the minimum number of males required to prevent extinction. Finally, we highlight the use of UAVs for assessing the mating dynamics of other marine, terrestrial or avian species, in which adults might exhibit visually detectable differences, such as sexual dimorphism, external body characteristics or grouping tendencies. This article is protected by copyright. All rights reserved.

Wide-ranging animals often traverse more than one country, making it important to establish international management co-operations and agreed protocols; however, accessing all available information on a given species, or even a population of interest, compiled by local, national and international organisations, is often complicated. In the case of sea turtles, this issue is further compounded because different life stages of the same population occupy different types of habitat; even as adults, while part of the population aggregates to breed at a single site in a given year, all other adult individuals are dispersed across foraging habitats up to 1000 km or more in distance. Information on the number of individuals, movement patterns and habitat use are needed to: (1) identify, select and conserve key breeding, foraging and developmental habitat effectively, (2) develop realistic models to predict current and future threat status of animals as accurately as possible, and (3) mitigate pressures operating in distant areas that, otherwise, might not be detected or linked to the population of interest. Here, I use sea turtles as a case study to show how our current knowledge on wide-ranging marine species is currently incomplete and, in many cases, disjointed. In particular, different techniques are often used to assimilate different types of information in different settings for different purposes (e.g. mark-recapture, genetics, strandings and nesting data). Ultimately, opening access to these data sources would facilitate major advances in research, as well as the transfer of knowledge and information to practitioners, allowing the effective implementation of conservation management.

Global biodiversity conservation relies on the efficient operation of protected areas, with revenue often originating from visitors. However, visitors generally select destinations where personal security and safety are guaranteed. Information about potential threats (natural hazards or terrorism) for global destinations is mainly released to the public via national safety travel advice and via global reports on disaster and terrorism risk indices. This study aimed to evaluate the extent to which different types of national security risks occur in countries that contain marine protected areas (MPAs) globally, towards highlighting the importance of incorporating these security issues into conservation management plans. Countries hosting most global marine protected areas (MPAs) have a low to medium risk of natural disasters. However, the analyses demonstrated that about one-third of MPAs are hosted by countries with lower income economies, which are also shown to have a lower capacity to cope with natural disasters. Of interest, countries with high terrorism risk host only a small fraction of global MPAs, with lower income countries being subjected to significantly higher terrorism risk than higher income countries. Overall, the results show that the chance of a country with an MPA being subject to a national security risk is generally low. However, countries with a higher risk of violence and natural disaster have lower coping capacities and weaker economies. Thus, MPAs in such countries should incorporate the risk of national security issues into their management schemes, particularly when visitor revenue is incorporated into wildlife protection and the employment of locals.

Often climatic niche models predict that any change in climatic conditions will impact species abundance or distribution. However, the accuracy of models that just incorporate climatic information to predict future species habitat use is widely debated. Alternatively, environmental conditions may simply need to be above some minimum threshold of climatic suitability, at which point, other factors drive population size. Using the example of nesting sites of loggerhead sea turtles (Caretta caretta) in the Mediterranean (n = 105), we developed climatic niche models to examine whether a climatic suitability threshold could be identified as a climatic indicator in order for large populations of a widespread species to exist. We then assessed the climatic suitability of sites above and below this threshold in the past (∼1900) and future (∼2100). Most large sites that are currently above the climatic threshold were above the threshold in the past and future, particularly when future nesting seasonality shifted to start 1–2 months earlier. Our analyses highlight the importance of future phenological shifts for maintaining suitability. Our results provide a positive outlook for sea turtle conservation, suggesting that climatic conditions may remain suitable in the future at sites that currently support large nesting populations. Our study also provides an alternative way of interpreting the outputs of climatic niche models, by generating a threshold as an index of a minimum climatic suitability required to sustain large populations. This type of approach offers the possibility to benefit from information provided by climate-driven models, while reducing their inherent uncertainties.

Sex-biased survival linked to anthropogenic threats places populations at risk. We show the utility of long-term multidecadal photo-identification (photo-id) combined with long-term high-resolution (Fastloc-GPS) satellite telemetry to investigate the links between mortality rates and patterns of movement for a wide-ranging, endangered marine vertebrate. Using a photo-identification database of 947 loggerhead turtles (Caretta caretta) compiled over 18 yr, we estimated greater annual survival rates of females (0.89; 95% confidence interval [CI] 0.87-0.90) compared to males (0.73; 95% CI 0.67-0.78). For males satellite-tracked across multiple breeding seasons, 100% (26 of 26) returned to the same breeding site, suggesting the calculated lower male survival rate was likely not due to emigration to breed elsewhere. 10,111 and 2,524 tracking days for males (n = 39 individuals) and females (n = 18 individuals), respectively, revealed different habitat-use patterns outside the breeding season: males tended to occupy foraging sites closer to shore and closer to breeding sites but, due to their generally annual breeding, compared to biennial breeding for females, males migrated further per year on average. These differences in movement patterns likely contribute to higher mortality in males through increased interaction with anthropogenic threats. Long-term identification coupled with tracking offers great promise for estimating the survival rates of other wide-ranging species.

We review how unmanned aerial vehicles (UAVs), often referred to as drones, are being deployed to study the abundance and behaviour of sea turtles, identifying some of the commonalities and differences with studies on other marine vertebrates, including marine mammals and fish. UAV studies of all three groups primarily focus on obtaining estimates of abundance, distribution and density, while some studies have provided novel insights on the body condition, movement and behaviour of individuals (including inter-specific interactions). We discuss the emerging possibilities of how UAVs can become part of the standard methodologies for sea turtle ecologists through combining information on abundance and behaviour. For instance, UAV surveys can reveal turtle densities and hence operational sex ratios of sea turtles, which could be linked to levels of multiple paternity. Furthermore, embedding UAV surveys within a mark-recapture framework will enable improved abundance estimates. The complexity of behaviours revealed by direct observations of sea turtles and animal-borne cameras can also be examined using UAV footage, complementing studies using electronic tags, such as time-depth recorders and satellite transmitters. Overall, UAVs provide a low-cost approach of quantifying the flexibility of marine animal behaviour, allowing us to integrate information on abundance to establish how individuals respond to the presence of other organisms and the immediate environment.

There have been efforts around the globe to track individuals of many marine species and assess their movements and distribution, with the putative goal of supporting their conservation and management. Determining whether, and how, tracking data have been successfully applied to address real-world conservation issues is, however, difficult. Here, we compile a broad range of case studies from diverse marine taxa to show how tracking data have helped inform conservation policy and management, including reductions in fisheries bycatch and vessel strikes, and the design and administration of marine protected areas and important habitats. Using these examples, we highlight pathways through which the past and future investment in collecting animal tracking data might be better used to achieve tangible conservation benefits.

Why females would mate with multiple partners and have multiple fathers for clutches or litters is a long-standing enigma. There is a broad dichotomy in hypotheses ranging from polyandry having benefits to simply being an unavoidable consequence of a high incidence of male-female encounters. If females simply give in to mating when it is too costly to avoid being harassed by males (convenience polyandry), then there should be a higher rate of mating as density increases. However, if females actively seek males because they benefit from multiple mating, then mating frequency, and consequently the incidence of multiple paternity of clutches, should be high throughout. To explore these competing explanations, here we review the incidence of multiple paternity for sea turtles nesting around the World. Across 30 rookeries, including all 7 species of sea turtle, the incidence of multiple paternity was only weakly linked to rookery size (r

Why females would mate with multiple partners and have multiple fathers for clutches or litters is a long-standing enigma. There is a broad dichotomy in hypotheses ranging from polyandry having benefits to simply being an unavoidable consequence of a high incidence of male-female encounters. If females simply give in to mating when it is too costly to avoid being harassed by males (convenience polyandry), then there should be a higher rate of mating as density increases. However, if females actively seek males because they benefit from multiple mating, then mating frequency, and consequently the incidence of multiple paternity of clutches, should be high throughout. To explore these competing explanations, here we review the incidence of multiple paternity for sea turtles nesting around the World. Across 30 rookeries, including all 7 species of sea turtle, the incidence of multiple paternity was only weakly linked to rookery size (r2=0.14). However, using high resolution at-sea GPS tracking we show that the specifics of movement patterns play a key role in driving packing density and hence the likely rate of male-female encounters. When individuals use the same focal areas, packing density could be 100× greater than when assuming individuals move independently. Once the extent of adult movements in the breeding season was considered so that movements and abundance could be combined to produce a measure of density, then across rookeries we found a very tight relationship (r2=0.96) between packing density and the incidence of multiple paternity. These findings suggest that multiple paternity in sea turtles may have no benefit, but is simply a consequence of the incidence of male-female encounters.

Here, we show how seasonal changes in animal density drive strategic shifts in the activities of wildlife-watching operators. These shifts result in high viewing intensity when animal densities are low, highlighting the need for modifications to existing wildlife-watching guidelines. We used the endangered loggerhead sea turtle Caretta caretta as a model species that exhibits staggered departure from an important breeding area (Zakynthos, Greece, Mediterranean) over a 2-month period (July to August) when tourism is at a peak, to investigate changes in wildlife-watching strategies, zoning effectiveness and voluntary guideline compliance over time. We used a combination of direct land-based observations, global positioning system tracking (of wildlife-watching vessels and turtles) and models. The modelled number of turtles present in the breeding area decreased from > 200 in July to

Many sea turtle populations are recovering, highlighting the value of conservation efforts and long-term monitoring., We document a tendency for published estimates of population size in sea turtles to be increasing rather than decreasing across the globe. To examine the population status of the seven species of sea turtle globally, we obtained 299 time series of annual nesting abundance with a total of 4417 annual estimates. The time series ranged in length from 6 to 47 years (mean, 16.2 years). When levels of abundance were summed within regional management units (RMUs) for each species, there were upward trends in 12 RMUs versus downward trends in 5 RMUs. This prevalence of more upward than downward trends was also evident in the individual time series, where we found 95 significant increases in abundance and 35 significant decreases. Adding to this encouraging news for sea turtle conservation, we show that even small sea turtle populations have the capacity to recover, that is, Allee effects appear unimportant. Positive trends in abundance are likely linked to the effective protection of eggs and nesting females, as well as reduced bycatch. However, conservation concerns remain, such as the decline in leatherback turtles in the Eastern and Western Pacific. Furthermore, we also show that, often, time series are too short to identify trends in abundance. Our findings highlight the importance of continued conservation and monitoring efforts that underpin this global conservation success story.

Determining the time of day that animals initiate and end migration, as well as variation in diel movement patterns during migration, provides insights into the types of strategy used to maximise energy efficiency and ensure successful completion of migration. However, obtaining this level of detail has been difficult for long-distance migratory marine species. Thus, we investigated whether the large volume of highly accurate locations obtained by Argos-linked Fastloc-GPS transmitters could be used to identify the time of day that adult green (n = 8 turtles, 9487 locations) and loggerhead (n = 46 turtles, 47,588 locations) sea turtles initiate and end migration, along with potential resting strategies during migration. We found that departure from and arrival at breeding, stopover and foraging sites consistently occurred during the daytime, which is consistent with previous findings suggesting that turtles might use solar visual cues for orientation. Only seven turtles made stopovers (of up to 6 days and all located close to the start or end of migration) during migration, possibly to rest and/or refuel; however, observations of day versus night speed of travel indicated that turtles might use other mechanisms to rest. For instance, turtles travelled 31% slower at night compared to day during their oceanic crossings. Furthermore, within the first 24 h of entering waters shallower than 100 m towards the end of migration, some individuals travelled 72% slower at night, repeating this behaviour intermittently (each time for a one-night duration at 3–6 day intervals) until reaching the foraging grounds. Thus, access to data-rich, highly accurate Argos-linked Fastloc-GPS provided information about differences in day versus night activity at different stages in migration, allowing us, for the first time, to compare the strategies used by a marine vertebrate with terrestrial land-based and flying species.

Although the number and extent of protected areas (PAs) are continuously increasing, their coverage of global biodiversity, as well as criteria and targets that underline their selection, warrants scrutiny. As a case study, we use a global dataset of sea turtle nesting sites (n = 2991) to determine the extent to which the existing global PA network encompasses nesting habitats (beaches) that are vital for the persistence of the seven sea turtle species. The majority of nesting sites (87%) are in the tropics, and are mainly hosted by developing countries. Developing countries contain 82% nesting sites, which provide lower protection coverage compared to developed countries. PAs encompass 25% of all nesting sites, of which 78% are in marine PAs. At present, most nesting sites in PAs with IUCN ratification receive high protection. We identified the countries that provide the highest and lowest nesting site protection coverage, and detected gaps in species-level protection effort within countries. No clear trend in protection coverage was found in relation to gross domestic product, the Global Peace Index or sea turtle regional management units; however, countries in crisis (civil unrest, war or natural catastrophes) provided slightly higher protection coverage of all countries. We conclude that global sea turtle resilience against threats spanning temperate to tropical regions require representative PA coverage at the species level within countries. This work is anticipated to function as a first step towards identifying specific countries or regions that should receive higher conservation interest by national and international bodies.

The establishment of protected corridors linking the breeding and foraging grounds of many migratory species remains deficient, particularly in the world’s oceans. For example, Australia has recently established a network of Commonwealth Marine Reserves, supplementing existing State reserves, to protect a wide range of resident and migratory marine species; however, the routes used by mobile species to access these sites are often unknown. The flatback marine turtle (Natator depressus) is endemic to the continental shelf of Australia, yet information is not available about how this species uses the marine area. We used a geospatial approach to delineate a coastal corridor from 73 adult female flatback postnesting migratory tracks from four rookeries along the north-west coast of Australia. A core corridor of 1,150 km length and 30,800 km2 area was defined, of which 52 % fell within 11 reserves, leaving 48 % (of equivalent size to several Commonwealth Reserves) of the corridor outside of the reserve network. Despite limited data being available for other marine wildlife in this region, humpback whale migratory tracks overlapped with 96 % of the core corridor, while the tracks of three other species overlapped by 5–10 % (blue whales, olive ridley turtles, whale sharks). The overlap in the distribution ranges of at least 20 other marine vertebrates (dugong, cetaceans, marine turtles, sea snakes, crocodiles, sharks) with the corridor also imply potential use. In conclusion, this study provides valuable information towards proposing new locations requiring protection, as well as identifying high-priority network linkages between existing marine protected areas.

Management strategies to protect endangered species primarily focus on safeguarding habitats currently perceived as important (due to high-density use, rarity or contribution to the biological cycle), rather than sites of future ecological importance. This discrepancy is particularly relevant for species inhabiting beaches and coastal areas that may be lost due to sea-level rise over the next 100 years through climate change. Here, we modelled four sea-level rise (SLR) scenarios (0.2, 0.6, 0.9 and 1.3 m) to determine the future vulnerability and viability of nesting habitat (six distinct nesting beaches totalling about 6 km in length) at a key loggerhead sea turtle (Caretta caretta) rookery (Zakynthos, Greece) in the Mediterranean. For each of the six nesting beaches, we identified (1) the area of beach currently used by turtles, (2) the area of the beach anticipated to become inundated under each SLR, (3) the area of beach anticipated to become unsuitable for nesting under each SLR, (4) the potential for habitat loss under the examined SLR, and (5) the extent to which the beaches may shift in relation to natural (i.e. cliffs) and artificial (i.e. beach front development) physical barriers. Even under the most conservative 0.2 m SLR scenario, about 38% (range: 31–48%) total nesting beach area would be lost, while an average 13% (range: 7–17%) current nesting beach area would be lost. About 4 km length of nesting habitat (representing 85% of nesting activity) would be lost under the 0.9 m scenario, because cliffs prevent landward beach migration. In comparison, while the other 2 km of beach (representing 15% nests) is also at high risk, it has the capacity for landward migration, because of an adjoining sand-dune system. Therefore, managers should strengthen actions on this latter area, as a climatically critical safeguard for future sea turtle nesting activity, in parallel to regularly assessing and revising measures on the current high-use nesting habitats of this important Mediterranean loggerhead population.

This study evaluates how key beach features influence suitability for nesting by Endangered loggerhead marine turtles Caretta caretta at an internationally important rookery on Zakynthos Island, Greece. During 2007–2009 we assimilated information on beach structure (elevation above sea level and width), the distribution of all nesting (turtle tracks that resulted in nests) and non-nesting (turtle tracks that did not result in nests) turtle emergences from the sea along 6 km of beach, nest placement parameters (distance from sea and elevation above sea level), and beach use by visitors. We found that turtles preferentially emerged on steeper sections of beach, with higher nesting densities occurring on the most environmentally stable beaches. Elevation was a more reliable indicator of nest placement (1 m above sea level) than distance to shore. However, because nests on steeper slopes are located closer to shore, the risk of damage by tourism is increased in such areas. We calculated a potential 36% overlap of natural nest locations with use of the beach by tourists; however, the recorded overlap was 7% because of existing management protocols. This overlap could be further reduced by focusing conservation effort (i.e. further restricting use by people) on beach sections with the steepest inclines. For example, slopes of > 22° comprise 1 km of total beach area annually, the closure of which (above the immediate shoreline to allow passage) would completely protect 50% of nests. This study shows the value of evidence-based management as a practical scientific tool to conserve threatened species in dynamic protected areas that are of both environmental and economic importance.

Marine Protected Areas (MPAs) now form an important part of marine conservation and fisheries management; hence, there is broad interest in developing procedures that optimize their design. We used data collected over a 10-year period (2003–2012) from direct surveys and >100 adult male and female loggerhead sea turtles (Caretta caretta) tracked with devices, including GPS loggers and Fastloc GPS-Argos, to consider the optimum design for a MPA at a globally important breeding area, where there is already an existing national marine park aiming to protect the population (Zakynthos, Greece). Turtles primarily used areas very close to shore (approx. 7 km in length by 1 km in width, within the

International audience; In 2010, an international group of 35 sea turtle researchers refined an initial list of more than 200 research questions into 20 metaquestions that were considered key for management and conservation of sea turtles. These were classified under 5 categories: reproductive biology, biogeography, population ecology, threats and conservation strategies. To obtain a picture of how research is being focused towards these key questions, we undertook a systematic review of the peer-reviewed literature (2014 and 2015) attributing papers to the original 20 questions. In total, we reviewed 605 articles in full and from these 355 (59%) were judged to substantively address the 20 key questions, with others focusing on basic science and monitoring. Progress to answering the 20 questions was not uniform, and there were biases regarding focal turtle species, geographic scope and publication outlet. Whilst it offers some meaningful indications as to effort, quantifying peer-reviewed literature output is obviously not the only, and possibly not the best, metric for understanding progress towards informing key conservation and management goals. Along with the literature review, an international group based on the original project consortium was assigned to critically summarise recent progress towards answering each of the 20 questions. We found that significant research is being expended towards global priorities for management and conservation of sea turtles. Although highly variable, there has been significant progress in all the key questions identified in 2010. Undertaking this critical review has highlighted that it may be timely to undertake one or more new prioritizing exercises. For this to have maximal benefit we make a range of recommendations for its execution. These include a far greater engagement with social sciences, widening the pool of contributors and focussing the questions, perhaps disaggregating ecology and conservation.

The long-term survival of species with temperature-dependant sex determination requires a sufficient range of incubation temperatures to ensure that both males and females are produced. The primary sex ratio of sea turtles is determined by the temperature experienced by eggs during the middle third of incubation. Here, we investigated the variability in the production of male and female offspring by loggerhead sea turtles Caretta caretta at six nesting beaches in the temperate breeding area of Zakynthos, Greece. Hatchling sex ratios were estimated using incubation durations and sand temperatures for 2007–2009, while the empirical relationship between air and sand temperature was used to infer historical (1875–2010) and future (2011–2100) hatchling sex ratios. First, all six beaches produced males; 55% of production was across five beaches, while 45% was focused on one beach (primarily in July). Second, male production varied across the season in different years; there was an initial peak in June, with production (rising, declining or plateauing) later in the season being regulated by August air temperatures. Interestingly, the annual male production rate estimated from the 3-year dataset (23%) was half that estimated from the 135-year reconstruction (50%), with the latter showing broad interannual variation. Finally, modelled predictions of future sex ratios ranged from a conservative 7.6% decline in male production by 2100 versus no production by 2038. This study provides a baseline from which predicted trends in hatchling sex ratio, in parallel to regular field assessments, could be used to provide guidelines for the selection of appropriate nest protection management protocols, to maintain required sex ratios that safeguard the future of this population.

Maintaining a high and stable body temperature is often critical for female ectotherms during reproduction. Yet this strategy may be energetically costly, and therefore challenging, during this period of already high-energy demand. 2. Here, the 6-week deployment of tri-axial accelerometers (n = 6) on a marine ectotherm, the loggerhead turtle (Caretta caretta), reproducing at the northern limit of the species' breeding range (i.e. in a thermally dynamic environment) revealed the behavioural mechanisms underly- ing its energy management strategy during the breeding season. 3. The estimated activity levels of female loggerheads using overall dynamic body acceleration (ODBA) were high during the breeding season, suggesting that marine turtles may not be able to remain inactive for long periods in the same manner as terrestrial ectotherms, because of the thermally dynamic nature of their environment. 4. However, activity levels were not constant throughout the season, being impacted by both ambient water temperature and female reproductive status. In cold water at the beginning of the nesting season, high levels of activity suggested that females behaviourally thermoregulated by seeking out warm water patches along the shoreline. Interactions with male turtles (courtship and ⁄ or avoidance) may also explain this high level of activity. As sea temperatures warmed up and the amount of energy devoted to reproduction probably increased, the turtles spent more time resting during long sequential flat-bottomed dives, and reduced any unnecessary locomotory activity. 5. Turtles may therefore adjust their activity patterns in response to seasonal variations in abi- otic (i.e. ambient temperature) and biotic (i.e. reproductive status) factors. This may help minimize activity-linked metabolic rate and maximize reproductive output over a season while breeding in thermally dynamic environments. 6. A mechanistic model gave support to these empirical results. The model revealed that actively maintaining high and stable body temperature is of clear benefit to female turtles at temperate breeding sites. While energetically costly, such active thermoregulatory behaviour may speed up egg maturation, allowing turtles to initiate nesting earlier in the season, and hence maximize reproductive output.

Aim Resources can shape patterns of habitat utilization. Recently a broad foraging dichotomy between oceanic and coastal sites has been revealed for loggerhead sea turtles (Caretta caretta). Since oceanic and coastal foraging sites differ in prey availability, we might expect a gross difference in home-range size across these habitats. We tested this hypothesis by equipping nine adult male loggerhead sea turtles with GPS tracking devices. Location National Marine Park of Zakynthos (NMPZ) Greece, central and eastern Mediterranean (Adriatic, Ionian and Aegean seas). Methods In 2007, 2008 and 2009, Fastloc GPS-Argos transmitters were attached to nine male loggerheads. In addition, a Sirtrack PTT unit was attached to one male in 2007. Four of the turtles were tracked on successive years. We filtered the GPS data to ensure comparable data volumes. Route consistency between breeding and foraging sites of the four re-tracked turtles was conducted. Foraging site home range areas and within site movement patterns were investigated by the fixed kernel density method. Results Foraging home range size ranged between circa 10 km2 at neritic habitats (coastal and open-sea on the continental shelf) to circa 1000 km2 at oceanic sites (using 90% kernel estimates), the latter most probably reflecting sparsely distributed oceanic prey. Across different years individuals did not follow exactly the same migration routes, but did show fidelity to their previous foraging sites, whether oceanic or neritic, with accurate homing in the final stages of migration. Main conclusions The broad distribution and diverse life-history strategies of this population could complicate the identification of priority marine protected areas beyond the core breeding site.

We assessed home range size for breeding loggerhead turtles ( Caretta caretta ) near the limit of the species range at the Greek island of Zakynthos in the Mediterranean. Thirteen adult females and seven adult males were tracked using GPS units (loggers and transmitters) during May and June of 2006, 2007 and 2008. Kernel analysis indicated that core home range sizes (50% estimator; range: 2.9–19.7 km 2 ) for both males and females were restricted to a 7.5 km tract of coastline. 15% of GPS locations fell outside of the national park protection zones, while within the protected breeding area 88% of GPS locations occurred in zones of minimal protection. Female home ranges were 64% larger in 2008 than in 2006 and 2007, indicating that several years monitoring may be required for the most effective designation of marine protected areas (MPAs). Ten of the tracked females departed the core breeding area on 15 occasions for periods of 1–15 days travelling distances of 10–100 km, although none nested at alternative breeding sites. The inter-annual variability of breeding area home range size and likelihood of incidence of forays appeared be correlated with barometric pressure. The movement responses of loggerheads to environmental conditions implicates an ability to switch nesting areas over small scales in response to climate change. However, such behaviour suggests the protection of existing core breeding sites may be inadequate, with policy makers being required to consider the protection of broader areas to encompass potential changes in the habitat needs of this species.