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Recording sand temperatures has become routine at many sea turtle nesting sites across the world given the impacts of incubation temperatures on hatchling sex ratios. However, the extent of thermal variability found at a nesting site has previously received little attention. Here we examine empirical sand temperature records across five atolls extending 250 km in the Chagos archipelago, Indian Ocean, between October 2012 and July 2023 and quantify the extent of spatial and temporal thermal variability. Our results suggest that sand temperatures at our study site vary seasonally and inter-annually, between beaches in the archipelago, and within beaches in different nesting habitats. The biggest drivers of thermal variability were seasonal and inter-annual differences, which modulated sand temperatures by up to 3.00 °C and 1.03 °C, respectively. Intra-beach and inter-beach variability further modulated temperatures by up to 0.56 °C and 0.85 °C, respectively. In addition, mean monthly sand temperatures were relatively low, suggesting that hatchling sex ratios are fairly balanced. The wide range of sand temperatures recorded at this nesting site suggests that it is likely both male-biased and female-biased clutches are produced during the nesting season. Quantifying thermal variability from a long-term sand temperature time series offers valuable insight into a population with temperature-dependent sex determination and, when possible, should be considered when modelling temperature impacts on hatchling sex ratios., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The importance of some ecosystems remains poorly understood. We showed that mesophotic ecosystems (30 to 150 m) are a key habitat for a critically endangered species, with strong evidence that a globally important population of adult hawksbill turtles ( Eretmochelys imbricata ) almost exclusively foraged at these depths on remote submerged banks. This discovery highlights the need for such areas to be included in conservation planning, for example, as part of the United Nations High Seas Treaty. We equipped nesting turtles with Fastloc-GPS (Global Positioning System) satellite tags at an Indian Ocean breeding area and they all traveled to deep foraging sites (6765 days of tracking data across 22 individuals including 183,921 dive-depth measurements) rather than shallow coral reef sites. Both chart depths and depth data relayed from the tags indicated that turtles foraged at mesophotic depths, the modal dive depths being between 35 and 40 m. We calculate that 55,554 km 2 of the western Indian Ocean alone consists of submerged banks between 30 and 60 m.

Climate warming and the feminization of populations due to temperature-dependent sex determination may threaten sea turtles with extinction. To identify sites of heightened risk, we examined sex ratio data and patterns of climate change over multiple decades for 64 nesting sites spread across the globe. Over the last 62 years the mean change in air temperature was 0.85°C per century (SD = 0.65°C, range = -0.53 to +2.5°C, n = 64 nesting sites). Temperatures increased at 40 of the 64 study sites. Female-skewed hatchling or juvenile sex ratios occurred at 57 of the 64 sites, with skews >90% female at 17 sites. We did not uncover a relationship between the extent of warming and sex ratio (r 62  = -0.03, p = .802, n = 64 nesting sites). Hence, our results suggest that female-hatchling sex ratio skews are not simply a consequence of recent warming but have likely persisted at some sites for many decades. So other factors aside from recent warming must drive these variations in sex ratios across nesting sites, such as variations in nesting behaviour (e.g. nest depth), substrate (e.g. sand albedo), shading available and rainfall patterns. While overall across sites recent warming is not linked to hatchling sex ratio, at some sites there is both is a high female skew and high warming, such as Raine Island (Australia; 99% female green turtles; 1.27°C warming per century), nesting beaches in Cyprus (97.1% female green turtles; 1.68°C warming per century) and in the Dutch Caribbean (St Eustatius; 91.5% female leatherback turtles; 1.15°C warming per century). These may be among the first sites where management intervention is needed to increase male production. Continued monitoring of sand temperatures and sex ratios are recommended to help identify when high incubation temperatures threaten population viability., (© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Projection models are being increasingly used to manage threatened taxa by estimating their responses to climate change. Sea turtles are particularly susceptible to climate change as they have temperature-dependent sex determination and increased sand temperatures on nesting beaches could result in the 'feminisation' of hatchling sex ratios for some populations. This study modelled likely long-term trends in sand temperatures and hatchling sex ratios at an equatorial nesting site for endangered green turtles (Chelonia mydas) and critically endangered hawksbill turtles (Eretmochelys imbricata). A total of 1078 days of sand temperature data were collected from 28 logger deployments at nest depth between 2018 and 2022 in Papua New Guinea (PNG). Long-term trends in sand temperature were generated from a model using air temperature as an environmental proxy. The influence of rainfall and seasonal variation on sand temperature was also investigated. Between 1960 and 2019, we estimated that sand temperature increased by ~0.6°C and the average hatchling sex ratio was relatively balanced (46.2% female, SD = 10.7). No trends were observed in historical rainfall anomalies and projections indicated no further changes to rainfall until 2100. Therefore, the sex ratio models were unlikely to be influenced by changing rainfall patterns. A relatively balanced sex ratio such as this is starkly different to the extremely female-skewed hatchling sex ratio (>99% female) reported for another Coral Sea nesting site, Raine Island (~850 km West). This PNG nesting site is likely rare in the global context, as it is less threatened by climate-induced feminisation. Although there is no current need for 'cooling' interventions, the mean projected sex ratios for 2020-2100 were estimated 76%-87% female, so future interventions may be required to increase male production. Our use of long-term sand temperature and rainfall trends has advanced our understanding of climate change impacts on sea turtles., (© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

How species respond to climate change may impact their extinction probability. Here we link climatology and ecology to tackle a globally important conservation question. For sea turtles, there are concerns that climate warming will cause both the feminization of populations as well as reduced hatchling survival. For 58 nesting sites across the world spanning all seven sea turtle species, we investigated whether warming might be avoided by shifts in nesting phenology to a cooler part of the year. We show that even with the most extreme phenological shift that has been reported to date-an 18-day advance in nesting per °C increase in sea surface temperature (SST)-temperatures will continue to increase at nesting sites with climate warming. We estimate that SST at nesting sites will rise by an average of 0.6°C (standard deviation = 0.9°C, n = 58) when we model a 1.5°C rise in SST combined with a best-case-scenario shift in nesting. Since sea turtles exhibit temperature-dependent sex determination, these temperature rises could lead to increasingly female-biased sex ratios as well as reduced hatchling production at sites across the world. These findings underscore concerns for the long-term survival of this iconic group., (© 2023 The Authors.)

Climate change is a clear and present threat to species survival. For species with temperature-dependent sex determination, including all sea turtles, it has been hypothesised that climate change may drive the creation of sex-ratio biases leading to population extinctions 1 . Through a global analysis across multiple species, we present the first direct empirical evidence for a demographic consequence of male scarcity in sea turtle populations, with a lower incidence of multiple paternity being found in populations with more extreme female-biased hatchling sex-ratio skews. For green turtles, when the female bias in hatchling sex ratio was >90%, the incidence of multiple paternity was low compared to other nesting sites, being 24.5% in the eastern Mediterranean (Cyprus), 36.4% on Redang Island (Malaysia) and 15.4% on the southern Great Barrier Reef (Heron Island, Australia) compared to higher values (range 61.1-91.7%) at other sites globally. These results suggest that a low incidence of multiple paternity may serve as a harbinger of future problems with egg fertility if males become even scarcer. Assessments of the incidence of multiple paternity at sites where adult males are expected to become scarce, such as Raine Island on the northern Great Barrier Reef in Australia, may help to identify when a lack of males raises the threat of local extinctions. In such cases, intervention to increase the production of male hatchlings may be needed., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2022 The Author(s). Published by Elsevier Inc. All rights reserved.)

The implications of logger accuracy and precision are rarely considered prior to their application in many ecological studies. We assessed the accuracy and precision of three temperature data loggers widely used in ecological studies (Hobo®, iButton® and TinyTag®). Accuracy was highest in TinyTags (95% of readings were within 0.23°C of the true temperature) and lowest in HOBOs and iButtons (95% of were readings within 0.43°C and 0.49°C of the true temperature, respectively). The precision (standard deviation of the repeat measurements) was greatest in TinyTags (0.04°C), followed by iButtons (0.17°C) and then HOBOs (0.22°C). As a case study, we then considered how modelled estimates of sea turtle hatchling sex ratios (derived from temperature), could vary as a function of logger accuracy. For example, at 29°C when the mean sex ratio derived was 0.47 female, the sex ratio estimate from a single logger could vary between 0.40 and 0.50 for TinyTags and 0.29 and 0.56 for both HOBOs and iButtons. Our results suggest that these temperature loggers can provide reliable descriptions of sand temperature if they are not over-interpreted. Logger accuracy must be considered in future ecological studies in which temperature thresholds are important.

We report Anthropogenic Marine Debris (AMD) in Chagos Archipelago in the Indian Ocean, globally amongst the most isolated island groups. AMD on 14 island beaches in five atolls were surveyed in 2019 using two techniques: Marine Debris Tracker (MDT) along littoral vegetation and photoquadrats in open beach. Over 60 % of AMD in both beach zones was composed of plastics, especially bottles and fragments (mean = 44.9 %, 27.2 %, range = 16.5-73.2 %, 4.8-55.9 % respectively in vegetation; mean = 28.7 %, 31.5 %, range = 17.7-40.7 %, 11.6-60.0 % respectively in open beach). The density of plastic debris in littoral vegetation (MDT data: 1995 bottles, 3328 fragments per 100 m 2 ) was 10-fold greater than in open beach (photoquadrat data: 184 bottles, 106 fragments per 100 m 2 ). Significant latitudinal variation in vegetation AMD occurred (8-fold greater in southern atolls, p = 0.006). AMD varied within island zones: most debris observed on oceanside beaches (oceanside vs lagoon, W = 365, p < 0.001; ocean vs island tip, W = 107, p = 0.034). Standardisation of surveys using the open-source MDT App is recommended. Debris accumulation hotspots overlapped with sea turtle nesting habitat, guiding future beach clean-up prioritisation., (Copyright © 2022 The Authors. Published by Elsevier Ltd.. All rights reserved.)

With some taxa, a reduction in the mean size of individuals may reflect over-harvesting and/or trophy hunting. However, we show that in sea turtles, a reduction in the mean size of breeding individuals may be part of the good news story of an expanding population. We describe a 70-fold increase in annual nest numbers on the island of Sal (Cape Verde, North Atlantic) between 2008 and 2020 (from 506 to 35 507 nests), making this now one of the largest loggerhead ( Caretta caretta ) nesting aggregations in the world. We use 20 128 measurements of the size of nesting turtles to show that their mean annual size has decreased by about 2.4 cm, from 83.2 to 80.8 cm. This decrease in the mean size of nesting turtles was not caused by the removal of larger turtles, for example by selective harvesting. Rather we develop a theoretical model to show than this decrease in mean size can be explained by an influx of first-time nesters, combined with a decrease in the size of those first-time nesters over time. A reduction in mean size of nesting turtles has been reported across the Atlantic, Pacific and Indian Oceans, and may be a common feature of population recoveries in sea turtles.

Development rate of ectothermic animals varies with temperature. Here we use data derived from laboratory constant temperature incubation experiments to formulate development rate models that can be used to model embryonic development rate in sea turtle nests. We then use a novel method for detecting the time of hatching to measure the in situ incubation period of sea turtle clutches to test the accuracy of our models in predicting the incubation period from nest temperature traces. We found that all our models overestimated the incubation period. We hypothesize three possible explanations which are not mutually exclusive for the mismatch between our modeling and empirically measured in situ incubation period: (1) a difference in the way the incubation period is calculated in laboratory data and in our field nests, (2) inaccuracies in the assumptions made by our models at high incubation temperatures where there is no empirical laboratory data, and (3) a tendency for development rate in laboratory experiments to be progressively slower as temperature decreases compared with in situ incubation., (© 2022 The Authors. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology published by Wiley Periodicals LLC.)

How animals navigate across the ocean to isolated targets remains perplexing greater than 150 years since this question was considered by Charles Darwin. To help solve this long-standing enigma, we considered the likely resolution of any map sense used in migration, based on the navigational performance across different scales (tens to thousands of kilometres). We assessed navigational performance using a unique high-resolution Fastloc-GPS tracking dataset for post-breeding hawksbill turtles ( Eretmochelys imbricata ) migrating relatively short distances to remote, isolated targets on submerged banks in the Indian Ocean. Individuals often followed circuitous paths (mean straightness index = 0.54, range 0.14-0.93, s.d. = 0.23, n = 22), when migrating short distances (mean beeline distance to target = 106 km, range 68.7-178.2 km). For example, one turtle travelled 1306.2 km when the beeline distance to the target was only 176.4 km. When off the beeline to their target, turtles sometimes corrected their course both in the open ocean and when encountering shallow water. Our results provide compelling evidence that hawksbill turtles only have a relatively crude map sense in the open ocean. The existence of widespread foraging and breeding areas on isolated oceanic sites points to target searching in the final stages of migration being common in sea turtles.

Reconstruction of past conditions provides important information on how ecosystems have been impacted by climate change, but generally for microhabitats worldwide there are no long-term empirical measurements. In these cases, there has been protracted debate about how various large-scale environmental proxies can best be used to reconstruct local temperatures. Here we help resolve this debate by examining how well environmental proxies hindcast sand temperatures at nest depths for five sea turtle nesting sites across the world. We link instrumental air temperature and sea surface temperature records with empirical sand temperature observations in the Atlantic (Ascension Island and Cape Verde), the Indian Ocean (Chagos Archipelago), the Caribbean (St Eustatius) and the Pacific (French Polynesia). We found strong correlations between sea surface temperatures, air temperatures and sand temperatures at all our study sites. Furthermore, Granger causality testing shows variations in sea surface temperature and air temperature precede variations in sand temperatures. We found that different proxies (air or sea temperature or a combination of both) predicted mean monthly sand temperatures within <0.5°C of empirical observations. Reconstructions of sand temperatures over the last 170 years reveal a slight warming of temperatures (maximum 0.5°C per century). An analysis of 36 published datasets revealed that the gradient of the relationship between sand temperature and air temperature is relatively constant, suggesting long-term changes in sand temperature could be extended around the world to include nesting sites where there are no empirical measurements of sand temperature. Our approaches are likely to have utility for a range of microhabitats where there is an interest in long-term changes in temperature., (© 2021 John Wiley & Sons Ltd.)