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Sea turtles are an iconic group of marine megafauna that have been exposed to multiple anthropogenic threats across their different life stages, especially in the past decades. This has resulted in population declines, and consequently many sea turtle populations are now classified as threatened or endangered globally. Although some populations of sea turtles worldwide are showing early signs of recovery, many still face fundamental threats. This is problematic since sea turtles have important ecological roles. To encourage informed conservation planning and direct future research, we surveyed experts to identify the key contemporary threats (climate change, direct take, fisheries, pollution, disease, predation, and coastal and marine development) faced by sea turtles. Using the survey results and current literature, we also outline knowledge gaps in our understanding of the impact of these threats and how targeted future research, often involving emerging technologies, could close those gaps.

Delineating spatial boundaries that accurately encompass complex, often cryptic, life histories of highly migratory marine megafauna can be a significant conservation challenge. For example, marine turtles range across vast ocean basins and coastal areas, thus complicating the evaluation of relative impacts of multiple overlapping threats and the creation of coherent conservation strategies. To address these challenges, spatially explicit ‘regional management units’ (RMUs) were developed in 2010 for all marine turtle species, globally. RMUs were intended to provide a consistent framework that organizes conspecific assemblages into units above the level of nesting rookeries and genetic stocks, but below the species level, within regional entities that may share demographic trajectories because they experience similar environmental conditions and other factors. From their initial conception, RMUs were intended to be periodically revised using new information about marine turtle distributions, life history, habitat use patterns, and population structure. Here, we describe the process used to update the 2010 RMU framework by incorporating newly published information and inputs from global marine turtle experts who are members of the IUCN Marine Turtle Specialist Group. A total of 48 RMUs for 6 of 7 marine turtle species and 166 distinct genetic stocks for all 7 species are presented herein. The updated RMU framework reflects a significant advance in knowledge of marine turtle biology and biogeography, and it provides improved clarity about the RMU concept and its potential applications. All RMU products have been made open access to support research and conservation initiatives worldwide.

Understanding the population genetic structure of the species is essential for determining the possible management units (UM) and their conservation and/or sustainable exploitation with it. Chelonia mydas is recognized as an endangered philopatric turtle. This work aims to describe the population structure of the green turtle in southwestern Cuba through traditional analytical approaches and allocation methods. The collections were made between 1998 and 2007 on five beaches in the Cuban southwest. Seven microsatellite loci from 149 individuals were amplified and genetic variability parameters were calculated. The population structure was inferred through the use of Wright's F, Analysis of Molecular Variance (AMOVA), and population assignment algorithms based on Bayesian analysis (STRUCTURE) and factorization of sparse non-negative matrices (implemented in R). Most of the loci were not in Hardy-Weinberg equilibrium, and several presented linkage disequilibrium. The AMOVA and differentiation statistics suggest the presence of structure at the geographical level analyzed. The highest value of ΔK and the lowest value of cross-entropy were reached for K = 2, a result that suggests that in southwestern Cuba there is the contribution of two ancestral populations of Chelonia mydas. Relative migration estimates indicate active genetic exchange between nesting colonies in southwestern Cuba. Received: 07.02.2022 Accepted: 09.12.2022 Editor: Erik García-Machado

La conservación efectiva de las Tortugas marinas depende del conocimiento de las interconexiones demográficas entre las poblaciones. Nosotros caracterizamos la variación genética dentro y entre dos colonias de anidación de tortuga verde (Chelonia mydas) en el suroeste de Cuba [N=28] y evaluamos estos datos en un contexto regional para comprender la contribución de cada colonia de anidación a la estructura genética de la especie, a partir de la secuenciación de 490 pb de la región de control del DNA mitocóndrico. En las colonias cubanas los haplotipos pertenecieron al mismo linaje (II) y el 71% de éstos resultaron endémicos, pero con baja representatividad. Entre las colonias de anidación cubanas no se encontró estructura genética significativa. Sin embargo, cuando se comparan los datos con las demás colonias del Gran Caribe se encontró estructura genética significativa (AMOVA), con mayor variación dentro que entre las colonias de anidación. La diferenciación genética (FST) estuvo positivamente correlacionada con la distancia geográfica entre las colonias de anidación. En la población cubana, un patrón de expansión súbita fue soportado por una distribución desigual observada. Sin embargo, El resultado de la Prueba de Fu rechazó el modelo estándar de neutralidad, probablemente como una consecuencia de migraciones recientes. Usando el análisis de clados anidados, se infirió un flujo de genes restringido y un aislamiento por distancia para los clados que contuvieron la mayoría de los haplotipos cubanos. Una fragmentación pasada y/ o una colonización a largas distancia fue inferida entre el total de clados anidados. A pesar de que la población de anidación del suroeste cubano comparte una conexión histórica con las otras poblaciones de la región, constituye una unidad demográfica separada que debe ser manejada independientemente.

Abstract Successful conservation of long‐lived species requires reliable understanding of long‐term trends and historical baselines. We present a framework for evaluating abundance trends and conservation outcomes for long‐lived marine species by integrating local ecological knowledge (LEK), ecological monitoring, and computer simulation, tested on a case‐study of long‐lived and heavily exploited green turtles (Chelonia mydas) in the Eastern Pacific. Models fit to LEK and monitoring data indicate that turtle abundance is increasing, but only after ∼40 years of safeguarding the species’ nesting and foraging habitats in Mexico. However, current abundance is at ∼60% of baseline levels and historic population structure has not been reestablished, indicating the need for sustained, long‐term conservation actions. We demonstrate the potential of linking LEK and ecological science to provide critical information for conservation, by establishing reference baselines and gauging population status with a long‐term historical perspective, while promoting equitable and sustainable futures.

Genetic monitoring of highly migratory endangered species is fundamental for effective management, particularly when they are shared internationally, and their populations need to be identified. A prime example is the green turtle, Chelonia mydas, whose genetic structure has been extensively studied in the Western Atlantic. Nevertheless, the identification of Cuban management units has remained uncertain, despite representing regionally significant nesting assemblages and occurring within a strategically central position. Compared to previous work, the current study used 800 bp mtDNA control region sequences and larger sample sizes (n = 189 from four nesting sites in SW Cuba). Of the 23 resolved haplotypes, nine were novel, fourteen were reported in Cuba for the first time, and eleven were endemic. Even though the distribution of nesting grounds barely spans 300 km, three management units were identified: Guanahacabibes-San Felipe (GUCB; with most of the endemic haplotypes), Isla de la Juventud (IJCB; with a predominance of haplotype CM-A13.1), and Cayo Largo (CLCB; with a haplotype profile closely related to Southern Caribbean rookeries). We discuss how the geographic distribution of mtDNA variation has likely been shaped by local and regional oceanic current patterns or derived from formerly hyperabundant regional populations. Genetic characterization of Cuban management units represents a significant contribution, filling critical knowledge gaps that have hampered the comprehensive mixed-stock analyses required to guide effective regional conservation strategies.

The olive ridley sea turtle is predominantly an epipelagic species with no apparent migration corridors. Research in feeding areas in other sea turtle species has identified aggregations of individuals from diverse backgrounds; however, no specific feeding areas have been identified for the olive ridley. We used mtDNA control region sequencing to identify the haplotype composition of 85 olive ridley turtles (adult and immature turtles from both sexes) captured and released in Central Mexican Pacific waters. Amplified fragments of the control region (751 bp) revealed the presence of 17 haplotypes (h = 0.5877 ± 0.0622 and π = 0.001698 ± 0.001185), and 16 of them were phylogenetically grouped within the lineage of the Eastern Pacific, but the haplotype Lo37 showed close evolutionary relationships with the lineage of the East Coast of India. Lagrangian drifter data showed that West–East transport from the Indo-Pacific to the tropical Eastern Pacific might be possible through passive drift. These results highlight the importance of more genetic studies offshore and the role of the ocean currents in the migratory behavior of olive ridleys, as well as the need to establish multinational strategies for the conservation and management of the species in international waters.

Deriving robust historical population trends for long-lived species subject to human exploitation is challenging in scenarios where long-term scientific data are scarce or unavailable, as often occurs for species affected by small-scale fisheries and subsistence hunting. The importance of Local Ecological Knowledge (LEK) in data-poor scenarios is increasingly recognized in conservation, both in terms of uncovering historical trends and for engaging community stewardship of historic information. Building on previous work in marine historical ecology and local ecological knowledge, we propose a mixed socio-ecological framework to reliably document and quantify LEK to reconstruct historical population trends. Our method can be adapted by interdisciplinary teams to study various long-lived taxa with a history of human use. We demonstrate the validity of our approach by reconstructing long-term abundance data for the heavily-exploited East Pacific green turtle (Chelonia mydas) in Baja California, Mexico, which was driven to near extinction by a largely unregulated fishery from the early 1950s to the 1980s. No scientific baseline abundance data were available for this time-frame because recent biological surveys started in 1995 after all green turtle fisheries in the area were closed. To fill this data gap, we documented LEK among local fishers using ethnographic methods and obtained verified, qualitative data to understand the socio-environmental complexity of the green turtle fishery. We then established an iterative framework to synthesize and quantify LEK using generalized linear models (GLMs) and nonlinear regression (NLR) to generate a standardized, LEK-derived catch-per-unit-effort (CPUE) time-series. CPUE is an index of abundance that is compatible with contemporary scientific survey data. We confirmed the accuracy of LEK-derived CPUE estimates via comparisons with fisheries statistics available for 1962–1982. We then modeled LEK-derived abundance trends prior to 1995 using NLR. Our model established baseline abundance and described historical declines, revealing that the most critical (exponential) decline occurred between 1960 and 1980. This robust integration of LEK data with ecological science is of critical value for conservation and management, as it contributes to a holistic view of a species’ historic and contemporary conservation status.

Temperature-dependent sex determination, or TSD, is a widespread phenomenon in reptiles. The shape of the relationship between constant incubation temperature and sex ratio defines the TSD pattern. The TSD pattern is considered a life-history parameter important for conservation because the wider the range of temperatures producing both sexes, the more resilient the species is to climate change impacts. We review the different published equations and methodologies that have been used to model TSD patterns. We describe a new flexible model that allows for an asymmetrical pattern around the pivotal temperature, which is the constant temperature producing both sexes in equal proportions. We show that Metropolis-Hastings with Markov chain produced by a Monte Carlo process has many advantages compared to maximum likelihood and is preferred. Finally, we apply the models to results from incubation experiments using eggs from the marine turtle Lepidochelys olivacea originating in Northeast Indian, East Pacific, and West Atlantic Regional Management Units (RMUs) and find large differences in pivotal temperatures but not in transitional ranges of temperatures.

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. [ABSTRACT FROM AUTHOR]

The nesting colony of green sea turtles Chelonia mydas at Guanahacabibes Peninsula Biosphere Reserve and National Park is one of the largest in the Cuban archipelago; however, little information about its nesting ecology is available. Temporal and spatial variation in nesting and reproductive success as well as morphometric characteristics of gravid females were used to ecologically characterize this colony. Nine beaches of the Southernmost coast of Guanahacabibes Peninsula were monitored for 14 years 1998-2012 to determine green turtle nesting activity, from May to September peak nesting season in this area. Beach dimensions were measured to determine nest density using the length and the area. Afterward the beaches were divided in two categories, index and secondary. Females were measured and tagged to compare new tagged females 823 with returning tagged females 140. Remigration interval was also determined. Temporal variation was identified as the annual number of nesting emergences and oviposits per female, with apparent peaks in reproductive activity on a biennial cycle in the first six years followed by periods of annual increase in nest number 2003-2008 and periods of decreasing number of nests 2010-2012. We also found intra-seasonal variation with the highest nesting activity in July, particularly in the second half of the month. The peak emergence time was 22:00-02:00hr. In terms of spatial variation, smaller beaches had the highest nest density and nesting was more frequent 6-9m from the high tide line, where hatchling production was maximized although hatchling success was high on average, above 80. Morphometric analysis of females was made and newly tagged turtles were smaller on average than remigrants. Our results are only a first attempt at characterizing Guanahacabibes populations but have great value for establishing conservation priorities within the context of national management plans, and for efficient monitoring and protection of nesting beaches.

Despite the long debate of whether or not multiple mating benefits the offspring, studies still show contradictory results. Multiple mating takes time and energy. Thus, if females fertilize their eggs with a single mating, why to mate more than once? We investigated and inferred paternal identity and number of sires in 12 clutches (240 hatchlings) of green turtles (Chelonia mydas) nests at Tortuguero, Costa Rica. Paternal alleles were inferred through comparison of maternal and hatchling genotypes, and indicated multiple paternity in at least 11 of the clutches (92%). The inferred average number of fathers was three (ranging from 1 to 5). Moreover, regression analyses were used to investigate for correlation of inferred clutch paternity with morphological traits of hatchlings fitness (emergence success, length, weight and crawling speed), the size of the mother, and an environmental variable (incubation temperature). We suggest and propose two different comparative approaches for evaluating morphological traits and clutch paternity, in order to infer greater offspring survival. First, clutches coded by the exact number of fathers and second by the exact paternal contribution (fathers who gives greater proportion of the offspring per nest). We found significant differences (P < 0.05) in clutches coded by the exact number of fathers for all morphological traits. A general tendency of higher values in offspring sired by two to three fathers was observed for the length and weight traits. However, emergence success and crawling speed showed different trends which unable us to reach any further conclusion. The second approach analysing the paternal contribution showed no significant difference (P > 0.05) for any of the traits. We conclude that multiple paternity does not provide any extra benefit in the morphological fitness traits or the survival of the offspring, when analysed following the proposed comparative statistical methods.

Previous genetic studies have demonstrated that natal homing shapes the stock structure of marine turtle nesting populations. However, widespread sharing of common haplotypes based on short segments of the mitochondrial control region often limits resolution of the demographic connectivity of populations. Recent studies employing longer control region sequences to resolve haplotype sharing have focused on regional assessments of genetic structure and phylogeography. Here we synthesize available control region sequences for loggerhead turtles from the Mediterranean Sea, Atlantic, and western Indian Ocean basins. These data represent six of the nine globally significant regional management units (RMUs) for the species and include novel sequence data from Brazil, Cape Verde, South Africa and Oman. Genetic tests of differentiation among 42 rookeries represented by short sequences (380 bp haplotypes from 3,486 samples) and 40 rookeries represented by long sequences (∼800 bp haplotypes from 3,434 samples) supported the distinction of the six RMUs analyzed as well as recognition of at least 18 demographically independent management units (MUs) with respect to female natal homing. A total of 59 haplotypes were resolved. These haplotypes belonged to two highly divergent global lineages, with haplogroup I represented primarily by CC-A1, CC-A4, and CC-A11 variants and haplogroup II represented by CC-A2 and derived variants. Geographic distribution patterns of haplogroup II haplotypes and the nested position of CC-A11.6 from Oman among the Atlantic haplotypes invoke recent colonization of the Indian Ocean from the Atlantic for both global lineages. The haplotypes we confirmed for western Indian Ocean RMUs allow reinterpretation of previous mixed stock analysis and further suggest that contemporary migratory connectivity between the Indian and Atlantic Oceans occurs on a broader scale than previously hypothesized. This study represents a valuable model for conducting comprehensive international cooperative data management and research in marine ecology.

Mitochondrial DNA analyses have been useful for resolving maternal lineages and migratory behavior to foraging grounds (FG) in sea turtles. However, little is known about source rookeries and haplotype composition of foraging green turtle aggregations in the southeastern Pacific. We used mitochondrial DNA control region sequences to identify the haplotype composition of 55 green turtles, Chelonia mydas, captured in foraging grounds of Gorgona National Park in the Colombian Pacific. Amplified fragments of the control region (457 bp) revealed the presence of seven haplotypes, with haplotype (h) and nucleotide (π) diversities of h = 0.300±0.080 and π = 0.009±0.005 respectively. The most common haplotype was CMP4 observed in 83% of individuals, followed by CMP22 (5%). The genetic composition of the Gorgona foraging population primarily comprised haplotypes that have been found at eastern Pacific rookeries including Mexico and the Galapagos, as well as haplotypes of unknown stock origin that likely originated from more distant western Pacific rookeries. Mixed stock analysis suggests that the Gorgona FG population is comprised mostly of animals from the Galapagos rookery (80%). Lagrangian drifter data showed that movement of turtles along the eastern Pacific coast and eastward from distant western and central Pacific sites was possible through passive drift. Our results highlight the importance of this protected area for conservation management of green turtles recruited from distant sites along the eastern Pacific Ocean.

Where conservation resources are limited and conservation targets are diverse, robust yet flexible priority-setting frameworks are vital. Priority-setting is especially important for geographically widespread species with distinct populations subject to multiple threats that operate on different spatial and temporal scales. Marine turtles are widely distributed and exhibit intra-specific variations in population sizes and trends, as well as reproduction and morphology. However, current global extinction risk assessment frameworks do not assess conservation status of spatially and biologically distinct marine turtle Regional Management Units (RMUs), and thus do not capture variations in population trends, impacts of threats, or necessary conservation actions across individual populations. To address this issue, we developed a new assessment framework that allowed us to evaluate, compare and organize marine turtle RMUs according to status and threats criteria. Because conservation priorities can vary widely (i.e. from avoiding imminent extinction to maintaining long-term monitoring efforts) we developed a "conservation priorities portfolio" system using categories of paired risk and threats scores for all RMUs (n = 58). We performed these assessments and rankings globally, by species, by ocean basin, and by recognized geopolitical bodies to identify patterns in risk, threats, and data gaps at different scales. This process resulted in characterization of risk and threats to all marine turtle RMUs, including identification of the world's 11 most endangered marine turtle RMUs based on highest risk and threats scores. This system also highlighted important gaps in available information that is crucial for accurate conservation assessments. Overall, this priority-setting framework can provide guidance for research and conservation priorities at multiple relevant scales, and should serve as a model for conservation status assessments and priority-setting for widespread, long-lived taxa.

BackgroundResolving threats to widely distributed marine megafauna requires definition of the geographic distributions of both the threats as well as the population unit(s) of interest. In turn, because individual threats can operate on varying spatial scales, their impacts can affect different segments of a population of the same species. Therefore, integration of multiple tools and techniques--including site-based monitoring, genetic analyses, mark-recapture studies and telemetry--can facilitate robust definitions of population segments at multiple biological and spatial scales to address different management and research challenges.Methodology/principal findingsTo address these issues for marine turtles, we collated all available studies on marine turtle biogeography, including nesting sites, population abundances and trends, population genetics, and satellite telemetry. We georeferenced this information to generate separate layers for nesting sites, genetic stocks, and core distributions of population segments of all marine turtle species. We then spatially integrated this information from fine- to coarse-spatial scales to develop nested envelope models, or Regional Management Units (RMUs), for marine turtles globally.Conclusions/significanceThe RMU framework is a solution to the challenge of how to organize marine turtles into units of protection above the level of nesting populations, but below the level of species, within regional entities that might be on independent evolutionary trajectories. Among many potential applications, RMUs provide a framework for identifying data gaps, assessing high diversity areas for multiple species and genetic stocks, and evaluating conservation status of marine turtles. Furthermore, RMUs allow for identification of geographic barriers to gene flow, and can provide valuable guidance to marine spatial planning initiatives that integrate spatial distributions of protected species and human activities. In addition, the RMU framework--including maps and supporting metadata--will be an iterative, user-driven tool made publicly available in an online application for comments, improvements, download and analysis.