Intra-Mediterranean migrations of loggerhead sea turtles (Caretta caretta) monitored by satellite telemetry

The movements of four Mediterranean loggerhead sea turtles (Caretta caretta; three females, one male) were tracked via satellite telemetry for between 108 and 457 days. Total length of the routes traveled by the turtles varied between 2554 and 7098 km, and the average travel rate was 1.2 km h^-1. Long-distance movement between the western and eastern Mediterranean basins followed a seasonal pattern and seemed to be triggered by temperature and food availability. In the autumn/winter months turtles moved from west to east in search of warmer waters, and returned to the western basin in spring, where food resources are generally more plentiful. Three (two females, one male) of the four turtles migrated eastward through the Straits of Messina, which is characterized by high fishing pressure and intense boat traffic. Information about turtle migration patterns and routes will serve to plan effective conservation strategies.     

Distribution of foraging habitats of male loggerhead turtles (Caretta caretta) as revealed by stable isotopes and satellite telemetry

Most studies on the foraging ecology of loggerhead turtles (Caretta caretta) have focused on adult females and juveniles. Little is known about the foraging patterns of adult male loggerheads. We analyzed tissues for carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) from 29 adult male loggerheads tracked with satellite transmitters from one breeding area in Florida, USA, to evaluate their foraging habitats in the Northwest Atlantic (NWA). Our study revealed large variations in δ¹³C and δ¹⁵N and a correlation between both δ¹³C and δ¹⁵N and the latitude to which the loggerheads traveled after the mating season, thus reflecting a geographic pattern in the isotopic signatures. Variation in δ¹³C and δ¹⁵N can be explained by differences in food web baseline isotopic signatures rather than differences in loggerhead trophic levels. Stable isotope analysis may help elucidate residency and migration patterns and identify foraging sea turtle subpopulations in the NWA due to the isotopically distinct habitats used by these highly migratory organisms.     

Long-term behavior at foraging sites of adult female loggerhead sea turtles (Caretta caretta) from three Florida rookeries

We used satellite telemetry to study behavior at foraging sites of 40 adult female loggerhead sea turtles (Caretta caretta) from three Florida (USA) rookeries. Foraging sites were located in four countries (USA, Mexico, the Bahamas, and Cuba). We were able to determine home range for 32 of the loggerheads. One turtle moved through several temporary residence areas, but the rest had a primary residence area in which they spent all or most of their time (usually >11 months per year). Twenty-four had a primary residence area that was <500 km² (mean = 191). Seven had a primary residence area that was ≥500 km² (range = 573–1,907). Primary residence areas were mostly restricted to depths <100 m. Loggerheads appeared to favor areas with larger-grained sediment (gravel and rock) over areas with smaller-grained sediment (mud). Short-term departures from primary residence areas were either looping excursions, typically involving 1–2 weeks of continuous travel, or movement to a secondary residence area where turtles spent 25–45 days before returning to their primary residence area. Ten turtles had a secondary residence area, and six used it as an overwintering site. For those six turtles, the primary residence area was in shallow water (<17 m) in the northern half of the Gulf of Mexico (GOM), and overwintering sites were farther offshore or farther south. We documented long winter dive times (>4 h) for the first time in the GOM. Characterizing behaviors at foraging sites helps inform and assess loggerhead recovery efforts.     

Magnetic orientation by hatchling loggerhead sea turtles (Caretta caretta) from the Gulf of Mexico

Loggerhead sea turtle hatchlings emerge from nests on either the east or west coast of the South Florida peninsula and then migrate offshore in opposite directions. Under laboratory conditions, magnetic cues induce east coast hatchlings to swim in directions that promote their transport by oceanic surface currents, such as the North Atlantic gyre. However, the surface currents used by west coast hatchlings are unknown. We examined the responses of west (Sarasota) hatchlings to magnetic cues in the Gulf of Mexico, the Florida Straits, and the Gulf Stream to determine their (1) likely migratory routes (2) orientation where currents lead into the Atlantic Ocean, and (3) orientation adjacent to Florida’s east coast. The results suggest that migration inside Gulf waters may be circuitous, that the turtles respond appropriately to enter Atlantic waters, and that orientation along Florida’s east coast probably promotes transport by the Gulf Stream into the North Atlantic gyre.     

Discrimination of ocean wave features by hatchling loggerhead sea turtles, Caretta caretta

At the beginning of their offshore migration, hatchling sea turtles orient directly into oceanic waves as they swim away from land. Recent experiments have demonstrated that hatchlings swimming underwater can determine the propagation direction of waves by monitoring the circular movements they experience as waves pass above. During July and August 1993, we studied how loggerhead sea turtle hatchlings (Caretta caretta L.) from the east coast of Florida, USA, responded to a range of wave parameters. We constructed a wave simulator to reproduce in air the circular movements that normally occur beneath small ocean waves. Hatchlings suspended in air and subjected to these orbital movements attempted to orient into simulated waves when periods and amplitudes were similar to those found near the Florida coast. Orbital movements with longer periods (greater than 10 s), however, failed to elicit responses. The results demonstrate that hatchling loggerheads can distinguish between waves with different periods and amplitudes, and that Florida hatchlings respond most strongly to orbital movements closely resembling those of waves that occur near their natal beach. Received: 28 May 1996 / Accepted: 17 September 1996     

Fine-scale distribution of juvenile Atlantic and Mediterranean loggerhead turtles (Caretta caretta) in the Mediterranean Sea

Loggerhead turtles nesting in the Mediterranean Sea exhibit remarkable genetic structuring. This paper tests the hypothesis that young loggerhead turtles from different rookeries do not distribute homogeneously among the major Mediterranean foraging grounds, due to a complex pattern of surface currents. We extracted long fragments of mitochondrial DNA from 275 stranded or bycaught juvenile turtles from six foraging grounds (Catalano-Balearic Sea, Algerian basin, Tyrrhenian Sea, Adriatic Sea, northern Ionian Sea and southern Levantine Sea). We used a Bayesian mixed-stock analysis to estimate the contributions from rookeries in the Mediterranean, the North-west Atlantic and Cape Verde to the studied foraging grounds. Differences were found in the relative contribution of juvenile turtles of Atlantic and Mediterranean origin to each foraging ground. A decreasing proportion of Atlantic juveniles was detected along the main surface current entering the Mediterranean, with a high prevalence of turtles from eastern Florida in the Algerian basin and lower numbers elsewhere. In regard to the turtles of Mediterranean origin, juveniles from Libya prevailed in central and western Mediterranean foraging grounds other than the Algerian basin. Conversely, the Adriatic Sea was characterised by a large presence of individuals from western Greece, while the southern Levantine Sea was inhabited by a heterogeneous mix of turtles from the eastern Mediterranean rookeries (Turkey, Lebanon and Israel). Overall, the distribution of juveniles may be related to surface circulation patterns in the Mediterranean and suggests that fisheries might have differential effects on each population depending on the overlap degree between foraging and fishing grounds.     

Correlation between stomach temperatures and ambient water temperatures in free-ranging loggerhead turtles,Caretta caretta

Instruments were attached to loggerhead turtles, Caretta caretta, at Kamoda Point, Japan in 1989 and on the Senri-coast, Japan in 1991. We simultaneously obtained stomach temperatures, ambient water temperatures and diving depths from four free-ranging loggerhead turtles during the internesting periods, using small recorders. These data were analyzed to understand how body temperature was maintained. Ambient water temperatures changed coincidentally when turtles moved up or down, but a stable stomach temperatures was maintained despite water temperature fluctuations of ca. 20 to 90 min in duration. Loggerhead turtles also experienced water temperature fluctuations longer than 24 h. Stomach temperatures responded to those water temperature changes with a time lag of several hours (160, 170, 230 and 240 min). Stomach temperatures were higher than ambient water temperatures throughout the experimental periods. The median values of the thermal difference between stomach and water temperatures were 1.1, 1.3, 1.5 and 1.7°C. The higher stomach temperatures are thought to be caused by metabolic heat production.     

Sex ratios of loggerhead sea turtles Caretta caretta during the juvenile pelagic stage

Sex ratios are a fundamental trait for species reproduction. In species with temperature-dependent sex determination (TSD), sex ratios are not necessarily even, which has important demographic consequences. We examined the sex ratio of juvenile pelagic stage loggerhead turtles Caretta caretta offshore Madeira Island, North Eastern Atlantic, using laparoscopy and histology. The overall sex ratio was 2:1 (F:M), significantly different from an even sex ratio. Although there was no apparent temporal variation, sex ratios among size classes were significantly different. The sex ratio of juveniles was compared with known sex ratios for the putative source rookery and found to be similar to the subadults’ sex ratio, but significantly less female-biased than the hatchlings sex ratio. This suggests overestimation of hatchlings sex ratios and/or, less likely, differential mortality of females during the first months of life. Alternatively, the Madeira Island aggregation may be recruiting males from other geographical sources such as the Mediterranean and the Cape Verde.     

Survival and remigration probabilities for loggerhead turtles (Caretta caretta) nesting in the eastern Gulf of Mexico

Few long-term mark-recapture tagging datasets exist to estimate population parameters for loggerhead sea turtle (Caretta caretta) recovery units. Using a two-state open robust design model, we analyzed a 20-year (1990–2009) mark-recapture dataset from the Keewaydin Island loggerhead nesting assemblage off the southwest coast of Florida (USA) in the eastern Gulf of Mexico. For this analysis, 2,292 turtle encounters were evaluated, representing 841 individual nesting turtles. Survival was estimated at 0.73 (95 % CI 0.69–0.76). This estimate is comparable with survival estimates elsewhere in the Peninsular Florida subpopulation and is among the lowest estimates for the Northwest Atlantic loggerhead population. We documented no changes in remigration rates or clutch frequency over time. These are the first survival and remigration probabilities estimated for a loggerhead nesting assemblage in the eastern Gulf of Mexico.     

Ecology of loggerhead marine turtles Caretta caretta in a neritic foraging habitat: movements, sex ratios and growth rates

Much is still to be learned about the spatial ecology of foraging marine turtles, especially for juveniles and adult males which have received comparatively little attention. Additionally, there is a paucity of ecological information on growth rates, size and age at maturity, and sex ratios at different life stages; data vital for successful population modelling. Here, we present results of a long-term (2002–2011) study on the movements, residency, growth and sex ratio of loggerhead turtles (Caretta caretta) in Amvrakikos Gulf (39°0′N 21°0′E), Greece, using satellite telemetry (N = 8) and ongoing capture–mark–recapture (CMR; N = 300 individuals). Individuals encountered at sea ranged from large juvenile to adult (46.2–91.5 cm straight carapace length) and demonstrated growth rates within published norms (<2.7 cm yr^?1) that slowed with increasing body size. We revealed that an unexpectedly high proportion of animals were male (>44 % of captures above 65 cm straight carapace length), compared to region-wide female-biased hatchling production, indicating sex-biased survival or possible behavioural drivers for likelihood of capture in the region. Satellite tracking confirmed that some turtles establish discrete, protracted periods of residency spanning more than 1 year, whilst others migrated away from the site. These findings are underlined by CMR results with individual capture histories spanning up to 7 years, and only 18 % of individuals being recaptured.     

Oceanographic influences on the dive behavior of juvenile loggerhead turtles (Caretta caretta) in the North Pacific Ocean

Satellite telemetry data from 17 juvenile loggerhead turtles (43.5–66.5 cm straight carapace length) were used in conjunction with oceanographic data to analyze the influence of regional and seasonal oceanography on dive behavior in the North Pacific Ocean. Combined dive behavior for all individuals showed that turtles spent more than 80% of their time at depths <5 m, and more than 90% of their time at depths <15 m. Multivariate classifications of dive data revealed four major dive types, three representing deeper, longer dives, and one representing shallower dives shorter in duration. Turtles exhibited variability in these dive types across oceanographic regions, with deeper, longer dives in the Hawaii longline swordfish fishing grounds during the first quarter of the year, as well as in the Kuroshio Extension Bifurcation Region and the region near the Baja California Peninsula, Mexico. Turtles in the Kuroshio Extension Bifurcation Region also exhibited dive variability associated with mesoscale eddy features, with turtles making deeper, longer dives while associated with the strongest total kinetic energy. Turtles in the central North Pacific exhibited seasonality in dive behavior that appeared to reflect synchronous latitudinal movements with the North Pacific Subtropical Front and the associated seasonal, large-scale oceanography. Turtles made deeper, longer dives during the first quarter of the year within this region, the reported time and area where the highest loggerhead bycatch occurs by the longline fishery. These results represent the first comprehensive study of dive data for this species in this region. The increased understanding of juvenile loggerhead dive behavior and the influences of oceanography on dive variability should provide further insight into why interactions with longline fisheries occur and suggest methods for reducing the bycatch of this threatened species.     

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.     

Foraging grounds, movement patterns and habitat connectivity of juvenile loggerhead turtles (Caretta caretta) tracked from the Adriatic Sea

Knowledge about migratory routes and highly frequented areas is a priority for sea turtle conservation, but the movement patterns of juveniles frequenting the Adriatic have not been investigated yet, although juveniles represent the bulk of populations. We tracked by satellite six juvenile and one adult female loggerhead from the north Adriatic. The results indicated that loggerhead juveniles (1) can either show a residential behaviour remaining in the Adriatic throughout the year or perform seasonal migrations to other areas, (2) can remain even in the coldest, northernmost area during winter, (3) can frequent relatively small foraging areas, (4) mostly frequent the eastern part of the Adriatic, and (5) follow preferred migratory routes along the western and eastern Adriatic coasts. The movements of the adult turtle also revealed (6) a behavioural polymorphism in Mediterranean adults, which included a lack of area fidelity and connection between distant neritic foraging grounds.     

Genetic structuring of immature loggerhead sea turtles (Caretta caretta) in the Mediterranean Sea reflects water circulation patterns

The analysis of mitochondrial DNA in loggerhead sea turtles (Caretta caretta) from eight foraging grounds in the Mediterranean and the adjoining Atlantic revealed deep genetic structuring within the western Mediterranean. As a consequence, the foraging grounds off the North-African coast and the Gimnesies Islands are shown to be inhabited mainly by turtles of the Atlantic stocks, whereas the foraging grounds off the European shore of the western Mediterranean are shown to be inhabited mainly by turtles from the eastern Mediterranean rookeries. This structuring is explained by the pattern of sea surface currents and water masses and suggests that immature loggerhead sea turtles entering the western Mediterranean from the Atlantic and the eastern Mediterranean remain linked to particular water masses, with a limited exchange of turtles between water masses. As the north of the western Mediterranean comprises mostly individuals from the highly endangered eastern Mediterranean rookeries, conservation plans should make it a priority to reduce the mortality caused by incidental by-catch in these areas.     

Body temperature independence of solar radiation in free-ranging loggerhead turtles,Caretta caretta,during internesting periods

Body temperatures, ambient water temperatures, light intensities and vertical positions (depth) of eight loggerhead turtles, Caretta caretta, were monitored by small recorders during internesting periods from 1991 through 1993 off Wakayama Prefecture, Japan. Body temperatures of eight loggerhead turtles were higher than ambient water temperatures through-out their internesting periods. Light intensities were compared with body temperatures and no evidence was obtained to suggest that the raised body temperatures were caused by the direct influence of solar radiation. Body temperatures were kept higher than water temperatures in cloudy weather or even at night. Mean thermal differences between body and water temperatures were significantly different among individuals, and larger turtles had a greater mean thermal difference. Elevations in body temperatures of adult loggerhead turtles can reasonably be assumed to result from the accumulation of metabolically produced heat. Surfacing times (spent at depths shallower than 2 m) of seven turtles were only 10.3 to 38.9% of their internesting periods, with the exception of one turtle who spent 66.3% of her time at the surface. Loggerhead turtles did not seem to bask positively at the sea surface to absorb radiative heat.     

Oceanic residents, neritic migrants: a possible mechanism underlying foraging dichotomy in adult female loggerhead turtles (Caretta caretta)

To reveal the mechanism underlying intrapopulation variation in the use of feeding habitats (oceanic vs. neritic) by adult female loggerhead turtles (Caretta caretta), we compared telomere length in the epidermis (a proxy for age) between oceanic- and neritic-foraging recruits (first-time nesters). Based on egg-yolk stable isotope ratios, recruits at Yakushima Island, Japan, were clearly divided into small oceanic planktivores and large neritic benthivores. There were no significant differences in telomere length between oceanic and neritic foragers, suggesting that they start reproduction at similar ages. Turtles that experienced faster growing conditions during their oceanic early lives may achieve sexual maturity there, while others may move from oceanic areas into neritic habitats, switching diets from nutrient-poor macroplankton to nutrient-rich benthic fauna in order to compensate for their earlier slow growth rate and continue their sexual development, reaching maturity in neritic waters.     

Development and pre-validation of a testosterone enzyme immunoassay (EIA) for predicting the sex ratio of immature loggerhead sea turtles (Caretta caretta) recovered along the western coast of the central Adriatic Sea

Plasma testosterone levels were evaluated, by validating and using a commercially available testosterone enzyme immunoassay (EIA) as an indicator of the sex of immature loggerhead turtles (Caretta caretta) recovered along the western coast of the central Adriatic Sea between November 2011 and February 2012. Testosterone levels were measured in blood samples collected from 28 immature turtles kept in short-term maintenance at the Fondazione Cetacea Rescue Centre (Italy). Overall, plasma testosterone ranged between 229.3 and 2628.6 pg ml^?1, suggesting that the EIA procedure is effective for determining androgen titers in immature loggerhead sea turtles. Analysis of the obtained data indicates an unbiased sex ratio supporting previous studies of juvenile loggerhead turtle sex ratios in the Mediterranean Sea. The present work can be considered a starting point for augmenting knowledge on the dynamics of juvenile loggerhead aggregations increasingly found in the northern-central Adriatic Sea and for promoting local management for conservation actions.