Global patterns of illegal marine turtle exploitation.

Human exploitation of wildlife for food, medicine, curios, aphrodisiacs, and spiritual artifacts represents a mounting 21st-century conservation challenge. Here, we provide the first global assessment of illegal marine turtle exploitation across multiple spatial scales (i.e., Regional Management Units [RMUs] and countries) by collating data from peer-reviewed studies, grey literature, archived media reports, and online questionnaires of in-country experts spanning the past three decades. Based on available information, we estimate that over 1.1 million marine turtles were exploited between 1990 and 2020 against existing laws prohibiting their use in 65 countries or territories and in 44 of the world's 58 marine turtle RMUs, with over 44,000 turtles exploited annually over the past decade. Exploitation across the 30-year period primarily consisted of green (56%) and hawksbill (39%) turtles when identified by species, with hawksbills (67%) and greens (81%) comprising the majority of turtles exploited in the 1990s and 2000s, respectively, and both species accounting for similar levels of exploitation in the 2010s. Although there were no clear overarching trends in the magnitude or spatial patterns of exploitation across the three decades, there was a 28% decrease in reported exploitation from the 2000s to the 2010s. The 10 RMUs with the highest exploitation in the 2010s included seven green and three hawksbill turtle RMUs, with most reported exploitation occurring in RMUs that typically exhibit a low risk of population decline or loss of genetic diversity. Over the past decade, the number of RMUs with "moderate" or "high" exploitation impact scores decreased. Our assessment suggests that illegal exploitation appears to have declined over the past decade and, with some exceptions, is primarily occurring in large, stable, and genetically diverse marine turtle populations.

Physiological determinants of the internesting interval in sea turtles: a novel 'water-limitation' hypothesis.

The internesting interval separates successive clutches of sea turtle eggs, and its duration varies both among and within species. Here, we review the potential physiological limits to this interval, and develop the hypothesis that desalination capacity limits the internesting interval owing to the requirement for water deposition in eggs. Sea turtles deposit 1-4 kg of water per clutch in egg albumen; for most species, this represents about 2% of adult body mass. We calculate how quickly turtles can recover this water by estimating maximal salt excretion rates, metabolic water production and urinary losses. From this water balance perspective, the 'water-limitation' hypothesis is plausible for green turtles but not for leatherbacks. Some plasma biochemistry studies indicate dehydration in sea turtles during the nesting season, although this is not a universal finding and these data have rarely been collected during the internesting interval itself. There is mixed support for a trade-off between clutch size and the length of the interval. We conclude that the 'water-limitation' hypothesis is plausible for most sea turtle species, but requires direct experimentation.

Potential limitations of behavioral plasticity and the role of egg relocation in climate change mitigation for a thermally sensitive endangered species.

Anthropogenic climate change is widely considered a major threat to global biodiversity, such that the ability of a species to adapt will determine its likelihood of survival. Egg-burying reptiles that exhibit temperature-dependent sex determination, such as critically endangered hawksbill turtles (Eretmochelys imbricata), are particularly vulnerable to changes in thermal regimes because nest temperatures affect offspring sex, fitness, and survival. It is unclear whether hawksbills possess sufficient behavioral plasticity of nesting traits (i.e., redistribution of nesting range, shift in nesting phenology, changes in nest-site selection, and adjustment of nest depth) to persist within their climatic niche or whether accelerated changes in thermal conditions of nesting beaches will outpace phenotypic adaption and require human intervention. For these reasons, we estimated sex ratios and physical condition of hatchling hawksbills under natural and manipulated conditions and generated and analyzed thermal profiles of hawksbill nest environments within highly threatened mangrove ecosystems at Bahía de Jiquilisco, El Salvador, and Estero Padre Ramos, Nicaragua. Hawksbill clutches protected in situ at both sites incubated at higher temperatures, yielded lower hatching success, produced a higher percentage of female hatchlings, and produced less fit offspring than clutches relocated to hatcheries. We detected cooler sand temperatures in woody vegetation (i.e., coastal forest and small-scale plantations of fruit trees) and hatcheries than in other monitored nest environments, with higher temperatures at the deeper depth. Our findings indicate that mangrove ecosystems present a number of biophysical (e.g., insular nesting beaches and shallow water table) and human-induced (e.g., physical barriers and deforestation) constraints that, when coupled with the unique life history of hawksbills in this region, may limit behavioral compensatory responses by the species to projected temperature increases at nesting beaches. We contend that egg relocation can contribute significantly to recovery efforts in a changing climate under appropriate circumstances.

Multiyear monitoring of survival following mitigation-driven translocation of a long-lived threatened reptile.

Translocation is used by managers to mitigate the negative impacts of development on species. Moving individuals to a new location is challenging, and many translocation attempts have failed. Robust, posttranslocation monitoring is therefore important for evaluating effects of translocation on target species. We evaluated the efficacy of a translocation designed to mitigate the effects of a utility-scale solar energy project on the U.S. federally listed Mojave desert tortoise (Gopherus agassizii). The species is a long-lived reptile threatened by a variety of factors, including habitat loss due to renewable energy development in the Mojave Desert and portions of the Colorado Desert in southern California (southwestern United States). We translocated 58 individual tortoises away from the project's construction site and intensively monitored them over 5 years (2012-2017). We monitored these individuals and tortoises located in the translocation release area (resident tortoises; n = 112) and control tortoises (n = 149) in a nearby location. We used our tortoise encounter data and known-fate survival models to estimate annual and cumulative survival. Translocated tortoises in each of 2 size classes (120-160 mm, >160 mm) did not survive at lower rates than resident and control tortoises over the study period. For models with different sets of biotic and abiotic covariates, annual and cumulative estimates of survival were always >0.87 and >0.56, respectively. Larger tortoises tended to have higher survival, but translocated tortoises were not differentially affected by the covariates used to model variation in survival. Based on these findings, our translocation design and study protocols could inform other translocation projects for desert species. Our case study highlights the benefits of combining rigorous scientific monitoring with well-designed, mitigation-driven management actions to reduce the negative effects of development on species of conservation concern.

Monitoreo Multianual de la Supervivencia de un Reptil Longevo en Peligro después de una Reubicación por Mitigación Resumen Los administradores utilizan la reubicación para mitigar los impactos negativos que el desarrollo tiene sobre las especies. El traslado de individuos hacia una nueva ubicación es todo un reto y muchos intentos de reubicación han fallado. Por esto el monitoreo robusto post-reubicación es importante para la evaluación de los efectos de la reubicación sobre las especies. Evaluamos la eficiencia de una reubicación diseñada para mitigar los efectos de un proyecto de energía solar fotovoltaica sobre la tortuga terrestre del desierto de Mojave (Gopherus agassizii), una especie en la lista federal estadunidense de especies en peligro. Los reptiles de esta especie son longevos y se encuentran en peligro por una variedad de factores, incluyendo la pérdida del hábitat por el desarrollo de energías renovables en el desierto de Mojave y en porciones del desierto del Colorado en el sur de California (suroeste de los Estados Unidos). Reubicamos a 58 individuos de esta especie para alejarlos del sitio de construcción del proyecto y los monitoreamos intensivamente durante cinco años (2012 - 2017). Monitoreamos a estos individuos y a las tortugas que ya se encontraban en el sitio de liberación (tortugas residentes; n = 112), así como a un grupo control de tortugas (n = 149) en una ubicación cercana. Usamos nuestros datos de encuentro con tortugas y modelos de supervivencia con destino conocido para estimar la supervivencia anual y acumulativa. Las tortugas reubicadas en cada una de las dos clases de tamaño (120-160 mm, >160 mm) no sobrevivieron a tasas más bajas que las residentes y las del grupo control durante el periodo de estudio. Para los modelos con conjuntos diferentes de co-variados bióticos y abióticos los estimados anuales y acumulativos de supervivencia fueron siempre >0.87 y >0.56, respectivamente. Las tortugas más grandes tendieron a tener una mayor supervivencia, aunque las tortugas reubicadas no se vieron afectadas diferencialmente por los co-variados que se usaron para modelar la variación de la supervivencia. Con base en estos hallazgos, nuestro diseño de reubicación y nuestros protocolos de estudio podrían informar a otros proyectos de reubicación para especies de desierto. Nuestro estudio de caso resalta los beneficios de la combinación del monitoreo científico riguroso con acciones de manejo bien diseñadas y llevadas por la mitigación para reducir los efectos negativos del desarrollo sobre las especies de importancia para la conservación.

Discrete, high-latitude foraging areas are important to energy budgets and population dynamics of migratory leatherback turtles.

Many broadly distributed migratory species exhibit fidelity to fine-scale areas that support vital life history requirements (e.g., resource acquisition, reproduction). Thus, such areas are critical for population dynamics and are of high conservation priority. Leatherback sea turtles are among the world's most widely distributed species, and their breeding and feeding areas are typically separated by thousands of kilometres. In this study, we analysed turtle-borne video data on daytime feeding rates and energy acquisition in Nova Scotia, Canada, to quantify the importance of this discrete, seasonal foraging area for leatherback energy requirements. Based on daytime foraging only, we estimate that a single foraging season in Nova Scotia could support 59% of a non-breeding leatherback's annual energy budget, and 29% of energetic requirements for a female on a typical 2-year reproductive cycle. However, maximum energy intake rates for leatherbacks are nearly four times lower than those of mammals and birds due the low energy content of leatherbacks' gelatinous zooplankton prey. These results illustrate that high quality, local-scale foraging areas such as Nova Scotia are critically important to the stability and future growth of the leatherback population in the Northwest Atlantic Ocean. Thus, as with other migratory species, efforts to reduce threats and maintain habitat quality in such areas should be high conservation priorities.

Protecting marine mammals, turtles, and birds by rebuilding global fisheries.

Reductions in global fishing pressure are needed to end overfishing of target species and maximize the value of fisheries. We ask whether such reductions would also be sufficient to protect non-target species threatened as bycatch. We compare changes in fishing pressure needed to maximize profits from 4713 target fish stocks-accounting for >75% of global catch-to changes in fishing pressure needed to reverse ongoing declines of 20 marine mammal, sea turtle, and seabird populations threatened as bycatch. We project that maximizing fishery profits would halt or reverse declines of approximately half of these threatened populations. Recovering the other populations would require substantially greater effort reductions or targeting improvements. Improving commercial fishery management could thus yield important collateral benefits for threatened bycatch species globally.

Mojave desert tortoise (Gopherus agassizii) thermal ecology and reproductive success along a rainfall cline.

Desert resource environments (e.g. microclimates, food) are tied to limited, highly localized rainfall regimes which generate microgeographic variation in the life histories of inhabitants. Typically, enhanced growth rates, reproduction and survivorship are observed in response to increased resource availability in a variety of desert plants and short-lived animals. We examined the thermal ecology and reproduction of US federally threatened Mojave desert tortoises (Gopherus agassizii), long-lived and large-bodied ectotherms, at opposite ends of a 250-m elevation-related rainfall cline within Ivanpah Valley in the eastern Mojave Desert, California, USA. Biophysical operative environments in both the upper-elevation, "Cima," and the lower-elevation, "Pumphouse," plots corresponded with daily and seasonal patterns of incident solar radiation. Cima received 22% more rainfall and contained greater perennial vegetative cover, which conferred 5°C-cooler daytime shaded temperatures. In a monitored average rainfall year, Cima tortoises had longer potential activity periods by up to several hours and greater ephemeral forage. Enhanced resource availability in Cima was associated with larger-bodied females producing larger eggs, while still producing the same number of eggs as Pumphouse females. However, reproductive success was lower in Cima because 90% of eggs were depredated versus 11% in Pumphouse, indicating that predatory interactions produced counter-gradient variation in reproductive success across the rainfall cline. Land-use impacts on deserts (e.g. solar energy generation) are increasing rapidly, and conservation strategies designed to protect and recover threatened desert inhabitants, such as desert tortoises, should incorporate these strong ecosystem-level responses to regional resource variation in assessments of habitat for prospective development and mitigation efforts.

Global patterns of marine mammal, seabird, and sea turtle bycatch reveal taxa-specific and cumulative megafauna hotspots.

Recent research on ocean health has found large predator abundance to be a key element of ocean condition. Fisheries can impact large predator abundance directly through targeted capture and indirectly through incidental capture of nontarget species or bycatch. However, measures of the global nature of bycatch are lacking for air-breathing megafauna. We fill this knowledge gap and present a synoptic global assessment of the distribution and intensity of bycatch of seabirds, marine mammals, and sea turtles based on empirical data from the three most commonly used types of fishing gears worldwide. We identify taxa-specific hotspots of bycatch intensity and find evidence of cumulative impacts across fishing fleets and gears. This global map of bycatch illustrates where data are particularly scarce--in coastal and small-scale fisheries and ocean regions that support developed industrial fisheries and millions of small-scale fishers--and identifies fishing areas where, given the evidence of cumulative hotspots across gear and taxa, traditional species or gear-specific bycatch management and mitigation efforts may be necessary but not sufficient. Given the global distribution of bycatch and the mitigation success achieved by some fleets, the reduction of air-breathing megafauna bycatch is both an urgent and achievable conservation priority.

Global priorities for marine biodiversity conservation.

In recent decades, many marine populations have experienced major declines in abundance, but we still know little about where management interventions may help protect the highest levels of marine biodiversity. We used modeled spatial distribution data for nearly 12,500 species to quantify global patterns of species richness and two measures of endemism. By combining these data with spatial information on cumulative human impacts, we identified priority areas where marine biodiversity is most and least impacted by human activities, both within Exclusive Economic Zones (EEZs) and Areas Beyond National Jurisdiction (ABNJ). Our analyses highlighted places that are both accepted priorities for marine conservation like the Coral Triangle, as well as less well-known locations in the southwest Indian Ocean, western Pacific Ocean, Arctic and Antarctic Oceans, and within semi-enclosed seas like the Mediterranean and Baltic Seas. Within highly impacted priority areas, climate and fishing were the biggest stressors. Although new priorities may arise as we continue to improve marine species range datasets, results from this work are an essential first step in guiding limited resources to regions where investment could best sustain marine biodiversity.

Resilience of marine turtle regional management units to climate change.

Enhancing species resilience to changing environmental conditions is often suggested as a climate change adaptation strategy. To effectively achieve this, it is necessary first to understand the factors that determine species resilience, and their relative importance in shaping the ability of species to adjust to the complexities of environmental change. This is an extremely challenging task because it requires comprehensive information on species traits. We explored the resilience of 58 marine turtle regional management units (RMUs) to climate change, encompassing all seven species of marine turtles worldwide. We used expert opinion from the IUCN-SSC Marine Turtle Specialist Group (n = 33 respondents) to develop a Resilience Index, which considered qualitative characteristics of each RMU (relative population size, rookery vulnerability, and genetic diversity) and non climate-related threats (fisheries, take, coastal development, and pollution/pathogens). Our expert panel perceived rookery vulnerability (the likelihood of functional rookeries becoming extirpated) and non climate-related threats as having the greatest influence on resilience of RMUs to climate change. We identified the world's 13 least resilient marine turtle RMUs to climate change, which are distributed within all three major ocean basins and include six of the world's seven species of marine turtle. Our study provides the first look at inter- and intra-species variation in resilience to climate change and highlights the need to devise metrics that measure resilience directly. We suggest that this approach can be widely used to help prioritize future actions that increase species resilience to climate change.

Shifting the life-history paradigm: discovery of novel habitat use by hawksbill turtles.

Adult hawksbill turtles (Eretmochelys imbricata) are typically described as open-coast, coral reef and hard substrate dwellers. Here, we report new satellite tracking data on female hawksbills from several countries in the eastern Pacific that revealed previously undocumented behaviour for adults of the species. In contrast to patterns of habitat use exhibited by their Caribbean and Indo-Pacific counterparts, eastern Pacific hawksbills generally occupied inshore estuaries, wherein they had strong associations with mangrove saltwater forests. The use of inshore habitats and affinities with mangrove saltwater forests presents a previously unknown life-history paradigm for adult hawksbill turtles and suggests a potentially unique evolutionary trajectory for the species. Our findings highlight the variability in life-history strategies that marine turtles and other wide-ranging marine wildlife may exhibit among ocean regions, and the importance of understanding such disparities from an ecological and management perspective.

Global conservation priorities for marine turtles.

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.

Regional management units for marine turtles: a novel framework for prioritizing conservation and research across multiple scales.

BACKGROUND: Resolving 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 FINDINGS: To 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/SIGNIFICANCE: The 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.

Conservation of marine megafauna through minimization of fisheries bycatch.

Many populations of marine megafauna, including seabirds, sea turtles, marine mammals, and elasmobranchs, have declined in recent decades due largely to anthropogenic mortality. To successfully conserve these long-lived animals, efforts must be prioritized according to feasibility and the degree to which they address threats with the highest relative impacts on population dynamics. Recently, Wilcox and Donlan (2007, Frontiers in Ecology and the Environment) and Donlan and Wilcox (2008, Biological Invasions) proposed a conservation strategy of "compensatory mitigation" in which fishing industries offset bycatch of seabirds and sea turtles by funding eradication of invasive mammalian predators from the terrestrial reproductive sites of these marine animals. Although this is a creative and conceptually compelling approach, we find it flawed as a conservation tool because it has narrow applicability among marine megafauna, it does not address the most pervasive threats to marine megafauna, and it is logistically and financially infeasible. Invasive predator eradication does not adequately offset the most pressing threat to most marine megafauna populations--fisheries bycatch. For seabird populations, fisheries bycatch and invasive predators infrequently are overlapping threats. Invasive predators have limited population-level impacts on sea turtles and marine mammals and no impacts on elasmobranchs, all of which are threatened by bycatch. Implementing compensatory mitigation in marine fisheries is unrealistic due to inadequate monitoring, control, and surveillance in the majority of fleets. Therefore, offsetting fisheries bycatch with eradication of invasive predators would be less likely to reverse population declines than reducing bycatch. We recommend that efforts to mitigate bycatch in marine capture fisheries should address multiple threats to sensitive bycatch species groups, but these efforts should first institute proven bycatch avoidance and reduction methods before considering compensatory mitigation.

Persistent leatherback turtle migrations present opportunities for conservation.

Effective transboundary conservation of highly migratory marine animals requires international management cooperation as well as clear scientific information about habitat use by these species. Populations of leatherback turtles (Dermochelys coriacea) in the eastern Pacific have declined by >90% during the past two decades, primarily due to unsustainable egg harvest and fisheries bycatch mortality. While research and conservation efforts on nesting beaches are ongoing, relatively little is known about this population of leatherbacks' oceanic habitat use and migration pathways. We present the largest multi-year (2004-2005, 2005-2006, and 2007) satellite tracking dataset (12,095 cumulative satellite tracking days) collected for leatherback turtles. Forty-six females were electronically tagged during three field seasons at Playa Grande, Costa Rica, the largest extant nesting colony in the eastern Pacific. After completing nesting, the turtles headed southward, traversing the dynamic equatorial currents with rapid, directed movements. In contrast to the highly varied dispersal patterns seen in many other sea turtle populations, leatherbacks from Playa Grande traveled within a persistent migration corridor from Costa Rica, past the equator, and into the South Pacific Gyre, a vast, low-energy, low-productivity region. We describe the predictable effects of ocean currents on a leatherback migration corridor and characterize long-distance movements by the turtles in the eastern South Pacific. These data from high seas habitats will also elucidate potential areas for mitigating fisheries bycatch interactions. These findings directly inform existing multinational conservation frameworks and provide immediate regions in the migration corridor where conservation can be implemented. We identify high seas locations for focusing future conservation efforts within the leatherback dispersal zone in the South Pacific Gyre.

Maternal investment in reproduction and its consequences in leatherback turtles.

Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001-2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.

Egg components, egg size, and hatchling size in leatherback turtles.

Relationships between egg size, egg components, and neonate size have been investigated across a wide range of oviparous taxa. Differences in egg traits among taxa reflect not only phylogenetic differences, but also interactions between biotic (i.e., maternal resource allocation) and abiotic (i.e. nest environment conditions) factors. We examined relationships between egg mass, egg composition, and hatchling size in leatherback turtles (Dermochelys coriacea) because of the unique egg and reproductive characteristics of this species and of sea turtles in general. Albumen comprised 63.0%+/-2.8% (mean+/-S.D.) of egg mass and explained most of the variation in egg mass, whereas yolk comprised only 33.0%+/-2.7%. Additionally, leatherback albumen dry mass was approximately 16% of albumen wet mass. Whereas hatchling mass increased significantly with egg mass (n = 218 clutches), hatchling mass increased by only approximately 2 g for each 10 g increase in egg mass and was approximately 10-20 g greater than yolk mass. Taken together, our results indicate that albumen might play a particularly significant role in leatherback embryonic development, and that leatherback eggs are both capable of water uptake from the nest substrate and also possess a large reservoir of water in the albumen. Relationships between egg mass and egg components, such as variation in egg mass being largely explained by variation in albumen mass and egg mass containing a relatively high proportion of albumen solids, are more similar to bird eggs than to eggs of other non-avian reptiles. However, hatchling mass correlates more with yolk mass than with albumen mass, unlike patterns observed in bird eggs of similar composition.

Bioenergetics and diving activity of internesting leatherback turtles Dermochelys coriacea at Parque Nacional Marino Las Baulas, Costa Rica.

Physiology, environment and life history demands interact to influence marine turtle bioenergetics and activity. However, metabolism and diving behavior of free-swimming marine turtles have not been measured simultaneously. Using doubly labeled water, we obtained the first field metabolic rates (FMRs; 0.20-0.74 W kg(-1)) and water fluxes (16-30% TBW day(-1), where TBW=total body water) for free-ranging marine turtles and combined these data with dive information from electronic archival tags to investigate the bioenergetics and diving activity of reproductive adult female leatherback turtles Dermochelys coriacea. Mean dive durations (7.8+/-2.4 min (+/-1 s.d.), bottom times (2.7+/-0.8 min), and percentage of time spent in water temperatures (Tw) < or =24 degrees C (9.5+/-5.7%) increased with increasing mean maximum dive depths (22.6+/-7.1 m; all P< or =0.001). The FMRs increased with longer mean dive durations, bottom times and surface intervals and increased time spent in Tw< or =24 degrees C (all r2> or =0.99). This suggests that low FMRs and activity levels, combined with shuttling between different water temperatures, could allow leatherbacks to avoid overheating while in warm tropical waters. Additionally, internesting leatherback dive durations were consistently shorter than aerobic dive limits calculated from our FMRs (11.7-44.3 min). Our results indicate that internesting female leatherbacks maintained low FMRs and activity levels, thereby spending relatively little energy while active at sea. Future studies should incorporate data on metabolic rate, dive patterns, water temperatures, and body temperatures to develop further the relationship between physiological and life history demands and marine turtle bioenergetics and activity.

Biotic and abiotic factors affect the nest environment of embryonic leatherback turtles, Dermochelys coriacea.

Clutches of leatherback turtles, Dermochelys coriacea, have lower hatching success than those of other sea turtles, but causes of high embryonic mortality are unknown. We measured characteristics of clutches along with spatial and temporal changes in PO(2) and temperature during incubation to determine the extent to which they affected the developmental environment of leatherback embryos. Minimum PO(2) in nests decreased as both the total number and mass of metabolizing embryos increased. Increases in both the number and mass of metabolizing embryos caused an increase in maximum nest temperature. However, neither PO(2) nor temperature was correlated with hatching success. Our measurements of relatively high nest PO(2) (lowest 17.1 kPa or 16.9% O(2)) indicate that hypoxia apparently does not cause the low hatching success of leatherback clutches. Oxygen partial pressure increased and temperature decreased from the center toward the periphery of leatherback nests. We inferred from these measurements that positions of eggs within nests vary in quality and potentially affect overall developmental success of entire clutches. The large metabolic mass of leatherback clutches and limits to gas flux imposed by the sand create a situation in which leatherback embryos collectively affect their own environment.