Inter-seasonal maintenance of individual nest site preferences in hawksbill sea turtles

Within a single population of hawksbill sea turtles (Eretmochelys imbricata), we found a behavioral polymorphism for maternal nest site choice with respect to beach microhabitat characteristics. Some females preferred to nest in littoral forest and in places with overstory vegetation cover, and others preferred to nest in more open, deforested areas. Nest site choice was consistent within and between nesting seasons two years apart. This was not a result of females simply returning to the same location along the shoreline; beach sections used by individual turtles varied between seasons. Nest site choice was not influenced by changes in beach environment (e.g., beach width and foliage cover) or by changes in females' reproductive output (e.g., clutch size), suggesting that fidelity to particular microhabitats is a major determinant of the observed nesting patterns. Because hawksbills exhibit temperature-dependent sex determination, if the behavioral polymorphism in nest site choice has a genetic basis, as is plausible, then this would have implications for sex ratio evolution and offspring survival. By taking an individual-based approach to the study of maternal behavior we reveal previously overlooked individual variation and hope to provide some impetus for more detailed studies of nest site choice.     

Influence of oceanic factors on long-distance movements of loggerhead sea turtles displaced in the southwest Indian Ocean

The routes of five satellite-tracked loggerhead turtles (Caretta caretta), subjected to an experimental translocation away from their usual migratory routes, have been analysed in relation to the concurrent oceanographic conditions. Remote sensing data on sea surface temperature and height anomalies, as well as trajectories of surface drifters were used, to get simultaneous information on the currents encountered by the turtles during their long-range oceanic movements. Turtles mostly turned out to move in the same direction as the main currents, and their routes were often influenced by circulation features they encountered. A comparison between turtle ground speeds with that of drifters shows that in several instances, the turtles did not drift passively with the currents but contributed actively to the overall movement. Two turtles embarked on an oceanic crossing, probably induced by seasonal changes in surface temperatures, a crossing that was largely determined by the main currents existing in the area.     

Strong site fidelity and longer internesting interval for solitary nesting olive ridley sea turtles in Brazil

Olive ridley sea turtles display two different types of nesting behavior: in arribada (synchronous mass nesting) or solitarily. Contrarily to arribadas, little has been published about solitary nesters. This study aimed to expand the knowledge on internesting interval and site fidelity of solitary nesting olive ridleys and to test a possible development of arribada nesting behavior. Data were collected in Sergipe (Brazil) over 125 km of beach from 10°30′S/36o23′W to 11°26′S/37o19′W, between nesting seasons 2004/2005 and 2006/2007. From 962 tagged females, 173 were seen renesting. The average internesting interval found was longer (22.35 ± 7.01 days) than previously described, which might relate to lower water temperatures during the internesting period. Olive ridleys at Sergipe showed high nesting site fidelity, with consecutive nesting events occurring in close proximity, non-randomly and dependently of previous events. Most of the consecutive nests were separated by 4.06–5.59 km. Development of arribada nesting behavior was not confirmed.     

Habitat use by immature loggerhead sea turtles in the Algerian Basin (western Mediterranean): swimming behaviour,seasonality and dispersal pattern

To study habitat use by loggerhead sea turtles in the Algerian Basin (western Mediterranean), ten juveniles (straight carapace length range: 39.0–63.3 cm) were tracked by satellite from March 2004 to September 2005. Swimming behaviour (characterized by speed of travel, time spent at the surface, and the cosine of turning angles) varied individually, but these differences were unrelated to body size. Despite individual differences in swimming behaviour, the ten immature loggerhead sea turtles spent most of their time in the oceanic waters of the Algerian Basin, although simulations indicated that the average tracking time (235.7 ± 98.7 SD days) was sufficiently long for them to leave the Algerian Basin and disperse through most of the Mediterranean. Furthermore, none of the ten turtles swam in any preferred direction, and their bearings were all randomly distributed. Finally, all them consistently avoided the continental shelf and did not migrate seasonally, as the average latitude, the average longitude, and the average distance of the population to the release point did not change seasonally. Seasonality also had only a weak influence in swimming behaviour, as the time spent at the surface during light hours was the only parameter that changed seasonally. We conclude that immature loggerhead sea turtles in the south of the western Mediterranean exhibit a strong fidelity to the Algerian Basin, where distribution is ruled mainly by the bathymetry, without any influence of seasonality. That fidelity to the Algerian Basin matches predictions based on genetic structuring and might result from a combination of factors: surface circulation patterns and habitat selection by the loggerhead sea turtles.     

Variation in adult annual survival probability and remigration intervals of sea turtles

We analyzed a large dataset to quantify adult annual survival probability and remigration intervals for the Tortuguero, Costa Rica green turtle population. Annual survival probability was estimated at 0.85 (95% CI 0.75–0.92) using a recovery model and at 0.85 (95% CI 0.83–0.87) using an open robust design model. The two most common modes of remigration are 2 and 3 years. Annual survival probability is lower and remigration intervals are shorter than for other green turtle populations. Explanations for short remigration intervals include reproductive compensation due to historic population declines, availability of better quality food items, favorable environmental conditions, and short distance to the main foraging grounds. Variation in survival and remigration intervals have profound consequences for management and life history evolution. The short remigration intervals of Tortuguero green turtles partly offset mortality caused by turtle fishing in Nicaragua and mean that low juvenile survival represents a more urgent threat to the population than low adult survival. Low adult survival probability could result in selective pressure for earlier age at maturity.     

Environmental pollution and biodiversity: Light pollution and sea turtles in the Caribbean

We examine the impact of pollution on biodiversity by studying the effect of coastal light pollution on the sea turtle population in the Caribbean. To this end we assemble a panel data set of sea turtle nesting activity and satellite-derived measures of nighttime light. Controlling for the surveyor effort, the local economic infrastructure, and spatial spillovers, we find that nighttime light significantly reduces the number of sea turtle nests. According to data on replacement costs for sea turtles raised in captivity, our result suggests that the increase in lighting over the last two decades has resulted in the loss of close to 1800 sea turtles in the Caribbean, worth up to $288 million. Incorporating our empirical estimates into a stage-structured population model, we discover that the dynamic effect of nighttime light on future generations of sea turtles is likely to be much larger, with a cost of approximately $2.8 billion for Guadeloupe alone. More generally, our study provides a new approach to valuing the cost of environmental pollution associated with species extinction.     

Incidental capture,direct mortality and delayed mortality of sea turtles in Australia's Northern Prawn Fishery

The species composition, catch and mortality rates of sea turtles captured incidentally by the tiger prawn fishery on Australia's northern coast in 1989 and 1990 were estimated by monitoring the fishery's catch. In 1990, the delayed rate of mortality from damage was estimated and the size composition was measured. Five species of turtles were captured: the flatback (Natator depressa, 59% of the total), loggerhead (Caretta caretta, 10%), olive ridley (Lepidochelys olivacea, 12%), green turtle (Chelonia mydas, 8%) and hawksbill (Eretmochelys imbricata, 5%). The turtle catches varied with water depth: the highest catch rates (0.068±0.006 turtles per trawl) were from trawls in water between 20 and 30 m deep, relatively few turtles (10%) were captured in water deeper than 40 m (25% of trawls). Catch rates varied with time of year: the highest catch rates were 0.098 (±0.013) turtles per trawl in winter. There was no significant difference in the overall catch rate (²= 0.047; p=0.8111; df=1) but a significant difference in mortality rate (²= 3.99; p<0.05; df=1) between the two years. The incidence of capture in the commercial fishery was 0.051 (±0.003) turtles per trawl towed for about 180 min, with 0.007 (±0.001) turtles per trawl drowning in the nets. There were no significant differences in the catch and mortality rates between the two years for any of the turtle species except the loggerhead, which had a significantly (² = 11.029; p=0.0013; df=1) lower catch rate in 1990 (0.002±0.001 turtles per trawl) than in 1989 (0.008±0.002 turtles per trawl), and a significantly higher mortality in 1990 (33%) than in 1989 (19%). Catch rates and mortality varied between the species: the flatback had the highest catch rate (0.030±0.002 turtles per trawl) but the lowest mortality (10.9%); the loggerhead had a catch rate of 0.005±0.001 turtles per trawl, and high mortality (21.9%); the olive ridley had a catch rate of 0.006±0.001 turtles per trawl and a low mortality (12.5%); the green turtle's catch rate was 0.004±0.001 per trawl and mortality 12.0%; the hawksbill had the lowest catch rate (0.002±0.001 turtles per trawl) but highest mortality (26.4%). Based on the fishing effort (27 049 d for 1989 and 25 746 d for 1990), we estimate that 5 503 (±424) turtles were caught and returned to the sea in 1989 and 5 238 (±404) in 1990, of which 567±140 drowned in 1989 and 943±187 in 1990. In 1990, an estimated 25% of all captured turtles suffered some non-lethal damage; an estimated 21% of turltes were captured comatose and 4% were injured. We conclude that, considering other threats, trawl-induced drowning is not the major impact on turtle populations in northern Australia, but that measures to reduce drowning and delayed mortality would be desirable.     

Impact of ocean acidification in the metabolism and swimming behavior of the dolphinfish (Coryphaena hippurus) early larvae

Since the industrial revolution, [CO₂]_atm has increased from 280 μatm to levels now exceeding 380 μatm and is expected to rise to 730–1,020 μatm by the end of this century. The consequent changes in the ocean’s chemistry (e.g., lower pH and availability of the carbonate ions) are expected to pose particular problems for marine organisms, especially in the more vulnerable early life stages. The aim of this study was to investigate how the future predictions of ocean acidification may compromise the metabolism and swimming capabilities of the recently hatched larvae of the tropical dolphinfish (Coryphaena hippurus). Here, we show that the future environmental hypercapnia (ΔpH 0.5; 0.16 % CO₂, ~1,600 μatm) significantly (p < 0.05) reduced oxygen consumption rate up to 17 %. Moreover, the swimming duration and orientation frequency also decreased with increasing pCO₂ (50 and 62.5 %, respectively). We argue that these hypercapnia-driven metabolic and locomotory challenges may potentially influence recruitment, dispersal success, and the population dynamics of this circumtropical oceanic top predator.     

Long-term and trans-life-cycle effects of exposure to ocean acidification in the green sea urchin Strongylocentrotus droebachiensis

Anthropogenic CO₂ emissions are acidifying the world’s oceans. A growing body of evidence demonstrates that ocean acidification can impact survival, growth, development and physiology of marine invertebrates. Here, we tested the impact of long-term (up to 16 months) and trans-life-cycle (adult, embryo/larvae and juvenile) exposure to elevated pCO₂ (1,200 μatm, compared to control 400 μatm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5-fold when acclimated to elevated pCO₂ for 4 months during reproductive conditioning, while no difference was observed in females acclimated for 16 months. Moreover, adult pre-exposure for 4 months to elevated pCO₂ had a direct negative impact on subsequent larval settlement success. Five to nine times fewer offspring reached the juvenile stage in cultures using gametes collected from adults previously acclimated to high pCO₂ for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO₂. pCO₂ had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO₂. These negative effects on settlement success and juvenile survival can be attributed to carry-over effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO₂ in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.     

The stomach contents of post-hatchling green and loggerhead sea turtles in the southwest Pacific: an insight into habitat association

Dietary information obtained from stomach contents can provide a wealth of information on an animal’s ecology. Where animals are cryptic, such as the post-hatchling life history stage of a sea turtle, the ecological insight that dietary analyses can provide, may be otherwise unobtainable. Investigations into post-hatchling turtle stomach contents have found planktonic organisms, dominated by pelagic molluscs and crustaceans, hydrozoans, Sargassum and fish eggs. The nature of these dietary organisms provides evidence for the widely accepted hypothesis that, with the exception of the flatback turtle (Natator depressus), the post-hatchling stage of a sea turtle’s life history is pelagic and oceanic. As the majority of studies that have investigated the stomach contents of post-hatchling sea turtles have been conducted on loggerhead turtles (Caretta caretta) in the northern Atlantic and Pacific Oceans, insight derived from dietary investigations into post-hatchling ecology is biased. This study investigates the diet of post-hatchling green turtles (Chelonia mydas) and loggerhead turtles in the southwest Pacific Ocean. Stomach contents were obtained from 55 green and loggerhead post-hatchling turtles that had stranded or been consumed by Coryphaena hippurus. Our findings demonstrate that loggerhead and green post-hatchlings in the southwest Pacific share similar feeding ecology and feed on a variety of neustonic items that are indicative of an oceanic and pelagic existence. The dietary items consumed by both species investigated belong to similar taxonomic groups as those found in previous studies with species level distinctions occurring owing to the different geographical location.     

Hitchhikers reveal cryptic host behavior: new insights from the association between Planes major and sea turtles in the Pacific Ocean

Studies that incorporate information from habitat-specific ecological interactions (e.g., epibiotic associations) can reveal valuable insights into the cryptic habitat-use patterns and behavior of marine vertebrates. Sea turtles, like other large, highly mobile marine vertebrates, are inherently difficult to study, and such information can inform the implementation of conservation measures. The presence of epipelagic epibionts, such as the flotsam crab Planes major, on sea turtles strongly suggests that neritic turtles have recently occupied epipelagic habitats (upper 200 m in areas with >200 m depth) and that epipelagic turtles spend time at or near the surface. We quantified the effects of turtle species, turtle size, and habitat (neritic or epipelagic) on the frequency of epibiosis (F ₀) by P. major on sea turtles in the Pacific Ocean. In neritic habitats, we found that loggerhead (F ₀ = 27.6 %) and olive ridley turtles (F ₀ = 26.2 %) host crabs frequently across a wide range of body sizes, and green turtles almost never host crabs (F ₀ = 0.7 %). These results suggest that loggerheads and olive ridleys display variable/flexible epipelagic-neritic transitions, while green turtles tend to transition unidirectionally at small body sizes. In epipelagic habitats, we found that loggerheads host crabs (F ₀ = 92.9 %) more frequently than olive ridleys (F ₀ = 50 %) and green turtles (F ₀ = 38.5 %). These results suggest that epipelagic loggerheads tend to spend more time at or near the surface than epipelagic olive ridleys and green turtles. Results of this study reveal new insights into habitat-use patterns and behavior of sea turtles and display how epibiont data can supplement data from more advanced technologies to gain a better understanding of the ecology of marine vertebrates during cryptic life stages.     

Primary site and initial products of ammonium assimilation in the symbiotic sea anemone Anemonia viridis

Invertebrates containing endosymbiotic dinoflagellate algae (zooxanthellae) retain excretory nitrogen, and many are able to take up ammonium from the surrounding seawater. However, the site of assimilation and role of nitrogen recycling between symbiont and host remains unclear. In the present study, ammonium uptake by the symbiotic sea anemone Anemonia viridis (Forskål) was examined by following the pathway of assimilation using ¹⁵N-enriched ammonium. Since zooxanthellae became enriched with ¹⁵N from ammonium at up to 17 times the rate of the host, they appear to be the primary site of assimilation. In the light, the rate of zooxanthellae enrichment at 20?M was twice that at 10?M, whereas the rate of host enrichment was not significantly affected by ammonium concentration. When anemones were incubated with [¹⁵N]ammonium in the dark, after 12?h without light the rate of enrichment was lowered in both zooxanthellae and host. However, while the enrichment of the host was significantly reduced when the light level was lowered from 300 to 150?μmol photons m^?2?s^?1, zooxanthellae enrichment was unchanged. Low molecular weight material from the zooxanthellae became enriched at 20 times the rate of that from the host, and enrichment was detected in the amino acids glutamate, glutamine, aspartate, alanine, glycine, phenylalanine, threonine, valine, tyrosine, and leucine from zooxanthellae. In the zooxanthellae, amino acids accounted for 65% of the total enrichment of low molecular weight material. Of the amino acids detected in zooxanthellae, over 90% of the enrichment was accounted for by glutamate, glutamine and aspartate. The enrichment of the amide group of glutamine was greater than that of the amine group of glutamate or glutamine, consistent with the glutamine synthetase/glutamine 2-oxoglutarate amidotransferase cycle as the mechanism of ammonium assimilation. To examine the flux of ¹⁵N from zooxanthellae to host, anemones were pulse-labelled with [¹⁵N]ammonium and then transferred to an unlabelled chase. Over a 2?h period there was no evidence for a flux of nitrogen from zooxanthellae to host. However, during the chase period, the enrichment of low molecular weight material declined and that of high molecular weight material increased in both zooxanthellae and host, indicating that protein was synthesized using ¹⁵N from ammonium in both components of the symbiosis. Again by using a pulse-chase system, it was found that glutamate was metabolised most rapidly by zooxanthellae, followed by (in order of decreasing rate of turnover) aspartate, alanine, glycine and valine (no data are available for glutamine). Unlike these amino acids, nitrogen was transferred to the essential amino acids phenylalanine and threonine, increasing their enrichment during the chase period. While recycled nitrogen is clearly important to this symbiosis, the mechanism by which it is cycled remains to be resolved.     

Temporal, spatial, and body size effects on growth rates of loggerhead sea turtles (Caretta caretta) in the Northwest Atlantic

In response to a call from the US National Research Council for research programs to combine their data to improve sea turtle population assessments, we analyzed somatic growth data for Northwest Atlantic (NWA) loggerhead sea turtles (Caretta caretta) from 10 research programs. We assessed growth dynamics over wide ranges of geography (9–33°N latitude), time (1978–2012), and body size (35.4–103.3 cm carapace length). Generalized additive models revealed significant spatial and temporal variation in growth rates and a significant decline in growth rates with increasing body size. Growth was more rapid in waters south of the USA (<24°N) than in USA waters. Growth dynamics in southern waters in the NWA need more study because sample size was small. Within USA waters, the significant spatial effect in growth rates of immature loggerheads did not exhibit a consistent latitudinal trend. Growth rates declined significantly from 1997 through 2007 and then leveled off or increased. During this same interval, annual nest counts in Florida declined by 43 % (Witherington et al. in Ecol Appl 19:30–54, 2009) before rebounding. Whether these simultaneous declines reflect responses in productivity to a common environmental change should be explored to determine whether somatic growth rates can help interpret population trends based on annual counts of nests or nesting females. Because of the significant spatial and temporal variation in growth rates, population models of NWA loggerheads should avoid employing growth data from restricted spatial or temporal coverage to calculate demographic metrics such as age at sexual maturity.     

Do mucilage events influence pico- and nanoplankton size and structure in the Adriatic sea?

Temporal changes in abundance and biomass of picophytoplankton, heterotrophic pico-eukaryotes, and nanoplankton assemblages were investigated along a transect crossing the Adriatic Sea, from the Italian to the Croatian coast. This 15-months (June 1999-August 2000) investigation allowed comparing microbial parameters during summer 1999 (year without mucilage) and summer 2000 when a major mucilage event occurred. Pico- and nanoplankton assemblages displayed significant differences between the 2 summer periods. The main differences can be summarized as: (i) presence of cyanobacterial blooms (up to 10⁸ cells l^-1) in summer 2000, absent in summer 1999; (ii) an increasing fraction of heterotrophic pico-eukaryotes (up to 5.0 × 10⁶ cells l^-1) and heterotrophic nanoplankton (size 2-5 µm) during mucilage event; (iii) a reduced abundance of small-sized (2-3 µm) phototrophic nanoplankton in summer 2000. Changes in community structure were signals of changes in trophic condition of the system, which resulted in a competitive advantage for small sized pico- and nanoheterotrophs. Data presented here indicated that mucilage events are associated with changes in microbial community structure and functioning in ambient water and induced the amplification of 3-step microbial food chain. The potential use of the heterotrophic pico-eukaryotes for describing alterations of the trophic pathways during mucilage events is discussed.     

Nest and maternal origin can influence morphology and locomotor performance of hatchling green turtles (Chelonia mydas) incubated in field nests

In numerous laboratory experiments involving the incubation of reptile eggs, both the maternal origin of eggs and the incubating environment (nest effect) have been demonstrated to influence hatchling phenotype. Although different hatchling phenotypes have been reported from natural nests, the separate effects of maternal origin and nest on hatchling phenotype in natural nests have not been demonstrated because in natural nests the two effects are confounded with each other. Here, we use a split clutch design to experimentally separate nest effects from maternal origin effects in field nests of green turtles (Chelonia mydas). We found both maternal origin and nest to influence hatchling morphology and locomotor performance in some but not all field nests. By using egg mass (maternal origin effect) and nest temperature (nest effect) in multiple regression analysis, we found maternal origin had a greater influence than nest temperature on the morphological attributes of hatchling mass and carapace size, but nest temperature had a greater influence than maternal origin on the performance attributes of self-righting time, self-righting propensity, swim thrust during the first 30 min of swimming, and power stroke rate during the first 30 min of swimming.     

Growth and content of energy reserves in juvenile sea scallops,Placopecten magellanicus,as a function of swimming frequency and water temperature in the laboratory

The effects of swimming frequency and water temperature on shell growth, tissue mass, and stored energy reserves of juvenile sea scallops, Placopecten magellanicus Gmelin, were examined in a factorial laboratory experiment spanning six weeks in July and August 1992. Individually tagged scallops of similar initial size (22.5±0.1 mm shell height, n=240) were induced to swim to exhaustion at three different swimming frequencies (every day, twice a week, or not at all) in two different water temperature regimes (4 to 7 or 7 to 13°C). The scallops were fed an ad libitum mixture of cultured microalgae. At the end of the experiment, cumulative increase in shell height, dry weight of soft tissues, condition index of dry adductor muscle (adductor muscle dry weight/soft tissue dry weight x 100) and total carbohydrate content of dry adductor muscle were measured for each scallop. Scallops at the higher temperature had significantly greater shell heights, and were in better metabolic condition as evidenced by significantly higher condition indices and muscle carbohydrate contents. The dry soft tissue weights did not differ significantly from their low temperature counterparts. Swimming frequency had no significant effect on shell height, dry tissue weight, or carbohydrate content, but condition index of the adductor muscle increased significantly with swimming frequency. These results show that not only was there no cumulative cost of swimming in terms of shell growth, total soft tissue weight, or carbohydrate content in young scallops, but that condition of adductor muscle tissue was higher in scallops that swam.     

Site fidelity and homing behavior in juvenile loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>)

To investigate site fidelity and homing behavior in juvenile loggerheads ( Caretta caretta, L.), a mark-recapture study spanning four years (1998–2001) was conducted in Core Sound, N.C., USA. Each year of the study, approximately half of the turtles captured were tagged and released near the capture sites ( n=207), while the remaining turtles were displaced 15–20 km and released ( n=198). Loggerheads in both groups were recaptured in equal proportions near the original capture sites and many individuals were also recaptured in subsequent years. These data imply that juvenile loggerheads often returned to their capture sites following displacement, because if turtles dispersed randomly or remained near their release sites, then fewer displaced turtles should have been caught again. Moreover, because turtles migrate out of North Carolina sounds each winter, turtles recaptured at the same locations in different years evidently returned to specific sites following long migrations. To further investigate homing behavior, a small number of displaced turtles ( n=28) were tracked using radio telemetry following their release. Although transmitters detached from most turtles within a few days, analyses of initial headings showed strong orientation in the direction of the capture site. In addition, four turtles successfully tracked for longer periods of time all returned rapidly to the vicinity of the capture location and remained in the area. Taken together, the results of this study indicate that juvenile loggerheads exhibit fidelity to specific areas during summer months and possess the navigational abilities to home to these areas following forced displacements and long-distance migrations.