Nest site fidelity of leatherback turtles at Playa Grande, Costa Rica

The possibility of nest site fidelity of leatherback turtles, Dermochelys coriacea, raises concerns about the merit of translocating poorly located clutches to a safe hatchery. Specifically, if some females habitually nest in areas where the probability of egg survival is low because of tidal inundation, nest relocation may inadvertently select for turtles that make poor nest site selections. We investigate the nest site fidelity of leatherbacks by studying consecutive nest site selections by leatherbacks at Playa Grande, Costa Rica, from 1993 to 2001. Along the ocean-to-vegetation axis, consecutive nest placements were independent (G₄²=4.737, P=0.315). Regardless of previous zone choices, leatherbacks typically nested in the open beach above the high tide line and away from vegetation. Thus, concerns about translocation of poorly located nests are unfounded. Along the coastal axis, however, an independence model was rejected (G₃₆²=145.179, P<0.001). The log-linear model-based analysis we developed and applied indicated that the pattern of dependence in nest site locations is more consistent with a hypothesis of spatial proximity in which leatherbacks simply nest close to the site of the previous nest than it is with a hypothesis of intrinsic fidelity to a specific location on the beach. Our model-based approach is suitable for other settings where longitudinal nesting data are available.     

Impact of nest relocation on the reproductive success of Loggerhead Turtles,Caretta caretta,in the Göksu Delta,Turkey (Reptilia: Cheloniidae)

When the nests of marine turtles are at a risk of inundation, relocation of the nests are often used in the conservation measures. Here, I determined the effect of nest relocation on Loggerhead Turtle (Caretta caretta) egg hatching success during the 2013 and 2014 nesting seasons in the Göksu Delta, Mersin, Turkey. I compared natural and relocated clutches, including those relocated before and after inundation, and evaluated 102 (94.6%) and 63 (81.1%) of survived nests in 2013 and 2014 respectively. Relocated nests experienced a 30% decrease in hatching success and a more prolonged incubation period compared to nests left in situ. Egg failure in nests relocated before and after inundation was similar in early-stage embryos, whereas it was three-fold higher in mid-stage embryos and two-fold lower in late-stage embryos. Thus, there was no significant difference in overall hatching success between the two relocation types. Moreover, there was no effect of delayed relocation of nests after inundation on hatching success. Possible impacts specific to the nesting site should be considered and explored before using nest relocation as a conservation tool. The relocation approach is recommended for nests at a high risk of inundation when the loss of nests is inevitable.     

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.     

On the dispersal of leatherback turtle hatchlings from Mesoamerican nesting beaches

So little is known about the early life history of leatherback turtles (Dermochelys coriacea) from hatchling to adulthood that this period has been termed the 'lost years'. For critically endangered eastern Pacific leatherback populations, continued and rapid declines underscore the urgent need to develop conservation strategies across all life stages. We investigate leatherback hatchling dispersal from four Mesoamerican nesting beaches using passive tracer experiments within a regional ocean modelling system. The evolution of tracer distribution from each of the nesting beaches showed the strong influence of eddy transport and coastal currents. Modelled hatchlings from Playa Grande, Costa Rica, were most likely to be entrained and transported offshore by large-scale eddies coincident with the peak leatherback nesting and hatchling emergence period. These eddies potentially serve as 'hatchling highways', providing a means of rapid offshore transport away from predation and a productive refuge within which newly hatched turtles can develop. We hypothesize that the most important leatherback nesting beach remaining in the eastern Pacific (Playa Grande) has been evolutionarily selected as an optimal nesting site owing to favourable ocean currents that enhance hatchling survival.     

Predaceous ants, beach replenishment, and nest placement by sea turtles

Ants known for attacking and killing hatchling birds and reptiles include the red imported fire ant (Solenopsis invicta Buren), tropical fire ant [Solenopsis geminata (Fabr.)], and little fire ant [Wasmannia auropunctata (Roger)]. We tested whether sea turtle nest placement influenced exposure to predaceous ants. In 2000 and 2001, we surveyed ants along a Florida beach where green turtles (Chelonia mydas L.), leatherbacks (Dermochelys coriacea Vandelli), and loggerheads (Caretta caretta L.) nest. Part of the beach was artificially replenished between our two surveys. As a result, mean beach width experienced by nesting turtles differed greatly between the two nesting seasons. We surveyed 1,548 sea turtle nests (2000: 909 nests; 2001: 639 nests) and found 22 ant species. S. invicta was by far the most common species (on 431 nests); S. geminata and W. auropunctata were uncommon (on 3 and 16 nests, respectively). In 2000, 62.5% of nests had ants present (35.9% with S. invicta), but in 2001, only 30.5% of the nests had ants present (16.4% with S. invicta). Turtle nests closer to dune vegetation had significantly greater exposure to ants. Differences in ant presence on turtle nests between years and among turtle species were closely related to differences in nest placement relative to dune vegetation. Beach replenishment significantly lowered exposure of nests to ants because on the wider beaches turtles nested farther from the dune vegetation. Selective pressures on nesting sea turtles are altered both by the presence of predaceous ants and the practice of beach replenishment.     

Comparative health assessment of Western Pacific leatherback turtles (Dermochelys coriacea) foraging off the coast of California, 2005-2007

Leatherback turtles (Dermochelys coriacea) are critically endangered, primarily threatened by the overharvesting of eggs, fisheries entanglement, and coastal development. The Pacific leatherback population has experienced a catastrophic decline over the past two decades. Leatherbacks foraging off the coast of California are part of a distinct Western Pacific breeding stock that nests on beaches in Indonesia, Papua New Guinea, and the Solomon Islands. Although it has been proposed that the rapid decline of Pacific leatherback turtles is due to increased adult mortality, little is known about the health of this population. Health assessments in leatherbacks have examined females on nesting beaches, which provides valuable biological information, but might have limited applicability to the population as a whole. During September 2005 and 2007, we conducted physical examinations on 19 foraging Pacific leatherback turtles and measured normal physiologic parameters, baseline hematologic and plasma biochemistry values, and exposure to heavy metals (cadmium, lead, and mercury), organochlorine contaminants, and domoic acid. We compared hematologic values of foraging Pacific leatherbacks with their nesting counterparts in Papua New Guinea (n=11) and with other nesting populations in the Eastern Pacific in Costa Rica (n=8) and in the Atlantic in St. Croix (n=12). This study provides the most comprehensive assessment to date of the health status of leatherbacks in the Pacific. We found significant differences in blood values between foraging and nesting leatherbacks, which suggests that health assessment studies conducted only on nesting females might not accurately represent the whole population. The establishment of baseline physiologic data and blood values for healthy foraging leatherback turtles, including males, provides valuable data for long-term health monitoring and comparative studies of this endangered population.     

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.     

Nest site selection in leatherbacks, Dermochelys coriacea: individual patterns and their consequences

We determined individual nest placement patterns for female leatherbacks nesting at Awa:la-Ya:lima:po, French Guiana, by measuring distance from the nest to several landscape features, such as the highest spring tide line (HSTL) and the vegetation line. Distance from the nest to the HSTL differed significantly between females, indicating the existence of individual nesting patterns. There was a significant repeatability of nest site choice relative to the HSTL, indicating that females showed within-individual consistency in their nest placement. Despite individual preferences, there was much within-individual variation and a lack of predictability in the nesting patterns; that is, the locations of subsequent nests could not be predicted based on knowledge of previous nest choices, indicating a certain degree of scatter. The significant repeatability suggests that nest choice behaviour in female leatherbacks is heritable and may show the potential for further evolution. We tested sea-finding ability of hatchlings, a potential consequence of nest site choice, in Matapica, Suriname, by using orientation arenas to quantify the strength and direction of travel after emergence. The orientation tests showed that hatchlings were unable to move seaward in vegetated arenas, providing evidence that vegetation acts as a strong selective pressure driving nest placement seaward. It appears that leatherbacks have adopted a regional rather than a local optimum for nest placement patterns, possibly resulting from their weak beach fidelity and the frequent erosion and destruction of their nesting beaches. We discuss the evolutionary and conservation implications for this species in the context of current environmental changes.     

Embryonic death is linked to maternal identity in the leatherback turtle (Dermochelys coriacea)

Leatherback turtles have an average global hatching success rate of ~50%, lower than other marine turtle species. Embryonic death has been linked to environmental factors such as precipitation and temperature, although, there is still a lot of variability that remains to be explained. We examined how nesting season, the time of nesting each season, the relative position of each clutch laid by each female each season, maternal identity and associated factors such as reproductive experience of the female (new nester versus remigrant) and period of egg retention between clutches (interclutch interval) affected hatching success and stage of embryonic death in failed eggs of leatherback turtles nesting at Playa Grande, Costa Rica. Data were collected during five nesting seasons from 2004/05 to 2008/09. Mean hatching success was 50.4%. Nesting season significantly influenced hatching success in addition to early and late stage embryonic death. Neither clutch position nor nesting time during the season had a significant affect on hatching success or the stage of embryonic death. Some leatherback females consistently produced nests with higher hatching success rates than others. Remigrant females arrived earlier to nest, produced more clutches and had higher rates of hatching success than new nesters. Reproductive experience did not affect stage of death or the duration of the interclutch interval. The length of interclutch interval had a significant affect on the proportion of eggs that failed in each clutch and the developmental stage they died at. Intrinsic factors such as maternal identity are playing a role in affecting embryonic death in the leatherback turtle.     

Density-dependent effects on hatching success of the olive ridley turtle, Lepidochelys olivacea

Historically, the olive ridley arribada at Playa Nancite, Costa Rica, was one of the largest olive ridley arribadas in the eastern Pacific with 70,000 nesting females in a year. Recently the Nancite arribada drastically declined. We hypothesized that the population decline at Playa Nancite could have been due to low hatching success as a result of the high density of nests on the beach, such that recruitment to the population was insufficient to balance losses. To test this hypothesis, we examined density-dependent effects on hatching success and their underlying mechanisms by experimentally manipulating nest densities in experimental plots on the nesting beach. We set up four nest-density treatments in five experimental blocks. We measured effects of density on hatching success, CO(2) and O(2) concentrations and temperature both within nests and in sand adjacent to nests frequently during incubation. Experimental nest densities affected hatching success with the highest density having the lowest hatching success. Higher nest density led to lower O(2) levels and higher CO(2) levels in the nest with greater changes in the latter part of the incubation. Highest temperatures occurred in high-density areas. Temperatures were lower in sand surrounding the nest than in the nest. Effects of density on temperature, CO(2) and O(2) were confirmed at a naturally high-density nesting beach, Playa La Flor, Nicaragua. Long-term failure in production of hatchlings due to historic high densities may have contributed to the decline of arribadas on Playa Nancite. Thus, density-dependent population control would have operated at the embryonic life stage in this population of olive ridley turtles.     

Beach dynamics and nest distribution of the leatherback turtle (Dermochelys coriacea) at Grande Riviere Beach, Trinidad & Tobago

Grande Riviere Beach in Trinidad and Tobago is an important nesting site in the Caribbean for the Critically Endangered leatherback sea turtle, Dermochelys coriacea. Community members were concerned that beach erosion and seasonal river flooding were destroying many of the nests deposited annually and thought that a hatchery was a possible solution. Over the 2001 turtle nesting season, the Institute of Marine Affairs (IMA) assessed the spatial and temporal distribution of nests using the Global Positioning System recorded to reference points, and beach dynamics using permanent bench mark profile stations, to determine areas of high risk and more stable areas for nesting. A total of 1449 leatherback nests were positioned. It was evident that at the start of the season in March, the majority of leatherback nests were deposited at the eastern section of the beach. After May, there was a continuing westward shift in nest distribution as the season progressed until August and beach erosion in the eastern section became predominant. The backshore remained relatively stable along the entire beach throughout the nesting season, and erosion was predominant in the foreshore at the eastern section of the beach, from the middle to the end of the season. Similar trends in accretion and erosion were observed in 2000. River flooding did not occur during the study period or in the previous year. With both high risk and more stable regions for turtle nesting available at Grande Riviere Beach, there was no compelling evidence to justify the need for a hatchery.     

Effects of Beach Nourishment on Sea Turtles: Review and Research Initiatives

Beach nourishment is an engineering solution to erosion of beaches. As in any restoration project, the goals of beach nourishment are the restoration of habitat to promote survival of plants and animals and to maintain aesthetically pleasing sites for humans. Unfortunately, beach nourishment sometimes alters parameters of the natural beach, decreasing the reproductive success of sea turtles. Engineers have recognized this problem and are working to improve nourishment practices. Biologists must specify problems incurred by sea turtles as a result of beach nourishment so that they may be addressed. A review of the literature on sea turtles and beach nourishment found certain problems repeatedly identified. For nesting females, characteristics induced by nourishment can cause (1) beach compaction, which can decrease nesting success, alter nest-chamber geometry, and alter nest concealment, and (2) escarpments, which can block turtles from reaching nesting areas. For eggs and hatchlings, nourishment can decrease survivorship and affect development by altering beach characteristics such as sand compaction, gaseous environment, hydric environment, contaminant levels, nutrient availability, and thermal environment. Also, nests can be covered with excess sand if nourishment is implemented in areas with incubating eggs. The extent and implication of each problem are discussed, and future research initiatives are proposed.     

An assessment of initial body size in loggerhead sea turtle (Caretta caretta) hatchlings in Turkey

Eggs, hatchlings, and adult loggerhead turtles, and incubation durations of clutches, were measured on three Turkish beaches (Dalyan, Fethiye and G?ksu Delta), and some physical features of nests were compared. These features were not statistically different among the beaches, except for nest depth and distance to the high water mark. There was a positive relationship between hatchling mass and egg size. The carapace length of hatchlings was correlated with both egg diameter and incubation duration. The duration of asynchronous emergence of hatchlings on Fethiye beach was slightly longer than on the other two beaches, and the size of hatchlings decreased as asynchronous emergence proceeded. Of the hatchlings that emerged first, those that died were significantly smaller in SCL and mass than those that lived. These results suggest that smaller hatchlings may not be vigorous enough to emerge earlier from nests, and that they may be less fit.     

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.     

Impact of Clutch Relocation on Green Turtle Offspring

ABSTRACT For species with temperature-dependent sex determination, such as marine turtles, global climate change poses numerous threats. At the nesting beach, rising temperatures are predicted to further skew already female-biased sex ratios and increase embryonic mortality; sea-level rise and resultant coastal squeeze may leave few alternative breeding habitats in developed regions. As a result, clutch relocation, a commonly used management tool to reduce egg loss, may become necessary for safeguarding populations. Although studies have examined the impact of relocation on clutch success, few have examined the impact of this practice on the sex or phenotypic characteristics of hatchlings produced. We used a randomized block design experiment to examine effects of relocation on green turtle (Chelonia mydas) clutches. We compared hatching success, thermal conditions, and size (length and mass) of hatchlings from in situ control clutches with those subjected to 2 relocation methods, while controlling for maternal and other environmental effects. Relocated clutches did not vary significantly from control clutches in incubation temperature or inferred sex ratios during the critical middle third of incubation when sex is thought to be determined. Hatchling size was also unaffected by relocation. Both relocation methods, however, resulted in a 20% reduction in hatching success in comparison to in situ clutches. Clutch relocation is, however, likely to affect the population primary sex ratio, when clutches are relocated from sites in proximity to the sea where tidal inundation is a threat. Here, cooler conditions are likely to produce more males than are the warmer female-producing temperatures higher up the beach. For clutches at risk, relocation is a viable process and does not appear to affect hatchling size or predicted sex ratios if relocation sites are selected in areas utilized by other females. We urge caution, however, when moving clutches from potentially male-producing sites, particularly given predicted impacts of climate change on already female-biased sex ratios.     

Nest Enlargement in Leaf-Cutting Ants: Relocated Brood and Fungus Trigger the Excavation of New Chambers

During colony growth, leaf-cutting ants enlarge their nests by excavating tunnels and chambers housing their fungus gardens and brood. Workers are expected to excavate new nest chambers at locations across the soil profile that offer suitable environmental conditions for brood and fungus rearing. It is an open question whether new chambers are excavated in advance, or will emerge around brood or fungus initially relocated to a suitable site in a previously-excavated tunnel. In the laboratory, we investigated the mechanisms underlying the excavation of new nest chambers in the leaf-cutting ant Acromyrmex lundi. Specifically, we asked whether workers relocate brood and fungus to suitable nest locations, and to what extent the relocated items trigger the excavation of a nest chamber and influence its shape. When brood and fungus were exposed to unfavorable environmental conditions, either low temperatures or low humidity, both were relocated, but ants clearly preferred to relocate the brood first. Workers relocated fungus to places containing brood, demonstrating that subsequent fungus relocation spatially follows the brood deposition. In addition, more ants aggregated at sites containing brood. When presented with a choice between two otherwise identical digging sites, but one containing brood, ants'' excavation activity was higher at this site, and the shape of the excavated cavity was more rounded and chamber-like. The presence of fungus also led to the excavation of rounder shapes, with higher excavation activity at the site that also contained brood. We argue that during colony growth, workers preferentially relocate brood to suitable locations along a tunnel, and that relocated brood spatially guides fungus relocation and leads to increased digging activity around them. We suggest that nest chambers are not excavated in advance, but emerge through a self-organized process resulting from the aggregation of workers and their density-dependent digging behavior around the relocated brood and fungus.     

Climate change and sea turtles: a 150-year reconstruction of incubation temperatures at a major marine turtle rookery

Sea turtles show temperature dependent sex determination. Using an empirical relationship between sand and air temperature, we reconstructed the nest temperatures since 1855 at Ascension Island, a major green turtle (Chelonia mydas) rookery. Our results show that inter‐beach thermal variations, previously ascribed to the albedo of the sand, which varies hugely from one beach to another, have persisted for the last century. Reconstructed nest temperatures varied by only 0.5 °C on individual beaches over the course of the nesting season, while the temperature difference between two key nesting beaches was always around 3 °C. Hence inter‐beach thermal variations are the main factor causing a large range of incubation temperatures at this rookery. There was a general warming trend for nests, with a mean increase in reconstructed nest temperatures for different months of between 0.36 and 0.49 °C for the last 100 years.     

The Effects of Artificial Beach Nourishment on Marine Turtles: Differences between Loggerhead and Green Turtles

Marine turtle reproductive success is correlated with the stability and quality of the nesting environment. Female marine turtles show fidelity to nesting beaches, making artificial beach nourishment practices directly relevant to their recovery. We evaluated the impacts of artificial beach nourishment on Loggerhead ( Caretta caretta ) and Green turtles ( Chelonia mydas ) between artificially nourished and nonnourished beaches. We observed reduced nesting success (ratio of nesting emergences to emergences not resulting in nest deposition) for both species. This negative effect lasted for one season in Loggerheads and for at least one season in Green turtles. Physical attributes of the fill sand did not impede nesting attempts. We argue that the decrease in nesting success resulted from an altered beach profile not favorable for nest deposition, which subsequently improved in later seasons as the beach equilibrated to a more natural slope. We observed a 52.2% decrease in reproductive output (hatchlings km⁻¹ yr⁻¹) for Loggerheads one year postnourishment, with a 44.1% increase observed the two seasons postnourishment. In Green turtles, a 0.8% reduction was observed the first season postnourishment, despite a 13% increase in the nonnourished area. The reduction in reproductive output in both cases was primarily a consequence of decreased nesting success, lowering nest numbers. These results reveal stronger negative effects of beach nourishment on Loggerheads compared to Green turtles and the importance of minimizing excessive nonnesting emergences associated with artificial beach nourishment. Nourished areas also experienced more than 600% increase in the number of Loggerhead hatchlings disoriented by artificial lighting over two years postnourishment.     

A potential tool to mitigate the impacts of climate change to the caribbean leatherback sea turtle

It is now well understood that climate change has the potential to dramatically affect biodiversity, with effects on spatio‐temporal distribution patterns, trophic relationships and survivorship. In the marine turtles, sex is determined by incubation temperature, such that warming temperatures could lead to a higher production of female hatchlings. By measuring nest temperature, and using a model to relate the incubation temperature to sex ratio, we estimate that Caribbean Colombian leatherback sea turtles currently produce approximately 92% female hatchlings. We modelled the relationship between incubation, sand and air temperature, and under all future climate change scenarios (0.4–6.0 °C warming over the next 100 years), complete feminization could occur, as soon as the next decade. However, male producing refugia exist in the periphery of smaller nests (0.7 °C cooler at the bottom than at the centre), within beaches (0.3 °C cooler in the vegetation line and inter‐tidal zone) and between beaches (0.4 °C higher on dark beaches), and these natural refugia could be assigned preferential conservation status. However, there exists a need to develop strategies that may ameliorate deleterious effects of climate‐induced temperature changes in the future. We experimentally shaded clutches using screening material, and found that it was effective in reducing nest temperature, producing a higher proportion of male hatchlings, without compromising the fitness or hatching success. Artificial shade in hatcheries is a very useful and simple tool in years or periods of high environmental temperatures. Nevertheless, this is only an emergency response to the severe impacts that will eventually have to be reversed if we are to guarantee the stability of the populations.     

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.