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IntroductionAn understanding of animal behavior is critical to determine their ecological role and to inform conservation efforts. However, observing hidden behaviors can be challenging, especially for animals that spend most of their time underwater. Animal-borne devices are valuable tools to estimate hidden behavioral states. MethodsWe investigated the fine-scale behavior of internesting hawksbill turtles using the mixed-membership method for movement (M4) which integrated dive variables with spatial components and estimated latent behavioral states. ResultsFive latent behavioral states were identified: 1) pre-nesting, 2) transit, 3) quiescence, and 4) area restricted search within and 5) near the residence of turtles. The last three states associated with a residency period, showed lower activity levels. Notably, when compared to other behaviors the pre-nesting exhibited shallower and remarkably long dives of up to 292 minutes. We noted high fidelity to residence core areas and nesting beaches, within and between nesting seasons, with residence areas decreasing within a season. DiscussionThe latent behaviors identified provide the most detailed breakdown of turtle movement behaviors during the internesting period to date, providing valuable insights into their ecology and behavior. This information can inform marine turtle conservation and management efforts since utilization distributions of individual behavioral states can be used to determine spatially-explicit susceptibility of turtles to various threats based on their behavior. The analyses of utilization distribution revealed a minimal overlap with existing marine protected areas (0.4%), and we show how a new proposal would expand protection to 30%. In short, this study provides valuable guidance for conservation and management of internesting marine turtles at a fine spatiotemporal resolution and can be used to enhance national action plans for endangered species, including the expansion of existing Marine Protected Areas. By flexibly incorporating biologically informative parameters, this approach can be used to study behavior outside of the hawksbill breeding season or even beyond this species. 

Abstract Species distribution models (SDMs) have become increasingly popular for making ecological inferences, as well as predictions to inform conservation and management. In predictive modeling, practitioners often use correlative SDMs that only evaluate a single spatial scale and do not account for differences in life stages. These modeling decisions may limit the performance of SDMs beyond the study region or sampling period. Given the increasing desire to develop transferable SDMs, a robust framework is necessary that can account for known challenges of model transferability. Here, we propose a comparative framework to develop transferable SDMs, which was tested using satellite telemetry data from green turtles (Chelonia mydas). This framework is characterized by a set of steps comparing among different models based on (1) model algorithm (e.g., generalized linear model vs. Gaussian process regression) and formulation (e.g., correlative model vs. hybrid model), (2) spatial scale, and (3) accounting for life stage. SDMs were fitted as resource selection functions and trained on data from the Gulf of Mexico with bathymetric depth, net primary productivity, and sea surface temperature as covariates. Independent validation datasets from Brazil and Qatar were used to assess model transferability. A correlative SDM using a hierarchical Gaussian process regression (HGPR) algorithm exhibited greater transferability than a hybrid SDM using HGPR, as well as correlative and hybrid forms of hierarchical generalized linear models. Additionally, models that evaluated habitat selection at the finest spatial scale and that did not account for life stage proved to be the most transferable in this study. The comparative framework presented here may be applied to a variety of species, ecological datasets (e.g., presence‐only, presence‐absence, mark‐recapture), and modeling frameworks (e.g., resource selection functions, step selection functions, occupancy models) to generate transferable predictions of species–habitat associations. We expect that SDM predictions resulting from this comparative framework will be more informative management tools and may be used to more accurately assess climate change impacts on a wide array of taxa. 

Abstract Small populations of imperiled species are susceptible to the negative consequences of skewed sex‐ratios. In imperiled species with environmental sex determination such as sea turtles, examining sex ratios across a range of environments and population abundance levels can provide insight into factors that influence population resilience, which can then be the foci of management plans for these species. Breeding sex ratios (the ratio of actively breeding males to females during a reproductive season; BSRs) extrapolated from genetic parentage analyses are a common approach for enumerating sex ratios in sea turtles. Such analyses also allow for the characterization of multiple paternity within sea turtle clutches, which should reflect BSRs and breeding behaviors. We characterized the first BSR for a breeding assemblage of loggerhead sea turtles (Caretta caretta) belonging to the temperate, low‐abundance Northern Gulf of Mexico Recovery Unit using genotypes of 16 microsatellite loci from nesting females and hatchlings. Unlike prior studies at both more‐tropical and more‐temperate, and higher‐abundance, Recovery Units in this region, we found a balanced BSR of 1.3:1 males:female and a low incidence (~17%) of multiple paternity. This suggests that there are relatively few males breeding at this assemblage and within this Recovery Unit. Beaches in this region are expected to produce substantial numbers of male hatchlings based on sand temperature data. The relative dearth of mature males may then be due to hydrologic disturbances that disproportionately affect the fitness and survival of male hatchlings, or due to demographic stochasticity. More work is needed to study the factors that might influence male hatchling production and fitness in this region, particularly as climate change is predicted to lead to feminization in global sea turtle populations. Our work demonstrates the broad utility of characterizing BSRs and other sex ratios across a range of populations in imperiled, environmentally sensitive species. 

Abstract Sea turtles are vulnerable to climate change since their reproductive output is influenced by incubating temperatures, with warmer temperatures causing lower hatching success and increased feminization of embryos. Their ability to cope with projected increases in ambient temperatures will depend on their capacity to adapt to shifts in climatic regimes. Here, we assessed the extent to which phenological shifts could mitigate impacts from increases in ambient temperatures (from 1.5 to 3°C in air temperatures and from 1.4 to 2.3°C in sea surface temperatures by 2100 at our sites) on four species of sea turtles, under a “middle of the road” scenario (SSP2‐4.5). Sand temperatures at sea turtle nesting sites are projected to increase from 0.58 to 4.17°C by 2100 and expected shifts in nesting of 26–43 days earlier will not be sufficient to maintain current incubation temperatures at 7 (29%) of our sites, hatching success rates at 10 (42%) of our sites, with current trends in hatchling sex ratio being able to be maintained at half of the sites. We also calculated the phenological shifts that would be required (both backward for an earlier shift in nesting and forward for a later shift) to keep up with present‐day incubation temperatures, hatching success rates, and sex ratios. The required shifts backward in nesting for incubation temperatures ranged from −20 to −191 days, whereas the required shifts forward ranged from +54 to +180 days. However, for half of the sites, no matter the shift the median incubation temperature will always be warmer than the 75th percentile of current ranges. Given that phenological shifts will not be able to ameliorate predicted changes in temperature, hatching success and sex ratio at most sites, turtles may need to use other adaptive responses and/or there is the need to enhance sea turtle resilience to climate warming. 

Satellite tracking has revolutionized our understanding of the behavior and ecology of sea turtles. However, most satellite tracking of sea turtles has been performed on breeding females, leaving knowledge gaps with regard to males. To inform future studies, we examined the peer-reviewed literature to summarize the contribution that satellite tracking has made to the spatial ecology of male turtles and describe how the published research varies across time, species, life stages, seasonal cycle phases, regions, and research topics. We systematically reviewed 61 publications reporting tracking of male sea turtles across 6 species. Loggerheads emerged as the most studied species, featuring in 49% of the publications, with green turtles and loggerheads having the highest number of tracked males. The North Atlantic was the most represented region (39% of publications), followed by the Mediterranean (29%). Most tracking of males has been undertaken at foraging areas (77%), often providing information on home range residency. This is followed by migration publications (44%), which have revealed differences among populations and between sexes, with the breeding period being the least studied. We highlight differences in the spatial ecology of males and females (e.g. residency in breeding areas and migration distance) and identified unanswered research questions about male sea turtles. To overcome the lack of knowledge about this important demographic group, efforts should be undertaken to increase the sample size and geographic coverage of tracked males, with special focus on flatback and Kemp’s ridleys, as well as studies in the Indian and Pacific Oceans. 

Marine protected areas (MPAs) are among the most widely used strategy to protect marine ecosystems and are typically designed to protect specific habitats rather than a single and/or multiple species. To inform the con- servation of species of conservation concern there is the need to assess whether existing and proposed MPA designs provide protection to these species. For this, information on species spatial distribution and exposure to threats is necessary. However, this information if often lacking, particularly for mobile migratory species, such as marine turtles. To highlight the importance of this information when designing MPAs and for assessments of their effectiveness, we identified high use areas of post-nesting hawksbill turtles (Eretmochelys imbricata) in Brazil as a case study and assessed the effectiveness of Brazilian MPAs to protect important habitat for this group based on exposure to threats. Most (88%) of high use areas were found to be exposed to threats (78% to artisanal fishery and 76.7% to marine traffic), where 88.1% were not protected by MPAs, for which 86% are exposed to threats. This mismatch is driven by a lack of explicit conservation goals and targets for turtles in MPA management plans, limited spatial information on species' distribution and threats, and a mismatch in the scale of conservation initiatives. To inform future assessments and design of MPAs for species of conservation concern we suggest that managers: clearly state and make their goals and targets tangible, consider ecological scales instead of political boundaries, and use adaptative management as new information become available. 

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 focused 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. 

Population abundance data are often used to define species’ conservation status. Abundance of marine turtles is typically estimated using nesting beach monitoring data such as nest counts and clutch frequency (CF, i.e., the number of nests female turtles lay within a nesting season). However, studies have shown that CF determined solely from nesting beach monitoring data can be underestimated, leading to inaccurate abundance estimates. To obtain reliable estimates of CF for hawksbill turtles in northeastern Brazil (6.273356° S, 35.036271° W), the region with the highest nesting density in the South Atlantic, data from beach monitoring and satellite telemetry were combined from 2014 to 2019. Beach monitoring data indicated the date of first nesting event, while state-space modeling of satellite telemetry data indicated the departure date of turtles, allowing calculations of residence length at breeding site and CF estimates based on internesting intervals. Females were estimated to nest up to six times within the nesting season with CF estimates between 4.5 and 4.8 clutches per female. CF estimates were used to determine the number of nesting females at the study site based in two approaches: considering and not considering transient turtles. Our approach and findings highlight that transients heavily influence CF estimates and need for reconsideration of how this key parameter is commonly determined for marine turtle populations and the use of beach monitoring data and satellite telemetry for estimations of CF