Different growth rates between loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) of Mediterranean and Atlantic origin in the Mediterranean Sea

We estimated for the first time the growth rates of loggerhead sea turtles of Mediterranean and of Atlantic origin found in the Mediterranean Sea, combining both skeletochronological and genetic analyses. Our growth models suggested that the growth rate of loggerhead sea turtles of Mediterranean origin was faster than that of their conspecifics with an Atlantic origin exploiting the feeding grounds in the Mediterranean Sea. The age at maturity for Mediterranean origin loggerhead sea turtles, estimated using our best fitting model, was 24 years, which suggests that loggerhead sea turtles nesting in the Mediterranean are not only smaller than those nesting in the western North Atlantic but also younger.     

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.     

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.     

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.     

Spatio-temporal distribution and migration of adult male loggerhead sea turtles (Caretta caretta) in the Mediterranean Sea: further evidence of the importance of neritic habitats off North Africa

Ten adult male loggerhead sea turtles, captured by trawlers or dip nets, were satellite-tracked from a neritic foraging ground in the Mediterranean in order to investigate adult spatio-temporal distribution and breeding migration. Five individuals migrated to potential breeding sites in Libya and one to Greece. The results complement previous studies and show that: (1) the Tunisian shelf may be more important for turtles from Libyan rookeries than previously thought; (2) male tracks corroborate a conservation hotspot previously identified for juveniles; (3) the north African coast represents a preferred migratory corridor, unless open sea routes are more direct; (4) adult males may exhibit high fidelity to relatively small areas, without evident seasonal differences; (5) adults home ranges were smaller and more neritic than juveniles frequenting the same area; (6) males may frequent multiple courtship areas; (7) the average remigration interval of males frequenting this region is longer than 1 year.     

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.     

Dune vegetation fertilization by nesting sea turtles

Sea turtle nesting presents a potential pathway to subsidize nutrient-poor dune ecosystems, which provide the nesting habitat for sea turtles. To assess whether this positive feedback between dune plants and turtle nests exists, we measured N concentration and delta15N values in dune soils, leaves from a common dune plant (sea oats [Uniola paniculata]), and addled eggs of loggerhead (Caretta caretta) and green turtles (Chelonia mydas) across a nesting gradient (200-1050 nests/km) along a 40.5-km stretch of beach in east central Florida, USA. The delta15N levels were higher in loggerhead than green turtle eggs, denoting the higher trophic level of loggerhead turtles. Soil N concentration and delta15N values were both positively correlated to turtle nest density. Sea oat leaf tissue delta15N was also positively correlated to nest density, indicating an increased use of augmented marine-based nutrient sources. Foliar N concentration was correlated with delta15N, suggesting that increased nutrient availability from this biogenic vector may enhance the vigor of dune vegetation, promoting dune stabilization and preserving sea turtle nesting habitat.     

Satellite tracking reveals a dichotomy in migration strategies among juvenile loggerhead turtles in the Northwest Atlantic

Few data are available on the movements and behavior of immature Atlantic loggerhead sea turtles (Caretta caretta) from their seasonal neritic foraging grounds within the western north Atlantic. These waters provide developmental habitat for loggerheads originating from several western Atlantic nesting stocks. We examined the long-term movements of 23 immature loggerheads (16 wild-caught and seven headstart turtles) characterizing their seasonal distribution, habitat use, site fidelity, and the oceanographic conditions encountered during their migrations. We identified two movement strategies: (1) a seasonal shelf-constrained north–south migratory pattern; and (2) a year-round oceanic dispersal strategy where turtles travel in the Gulf Stream to the North Atlantic and their northern dispersal is limited by the 10–15°C isotherm. When sea surface temperatures dropped below 20°C, neritic turtles began a migration south of Cape Hatteras, North Carolina (USA) where they established fidelity to the waters between North Carolina’s Outer Banks and the western edge of the Gulf Stream along outer continental shelf. Two turtles traveled as far south as Florida. Several turtles returned to their seasonal foraging grounds during subsequent summers. Northern movements were associated with both increased sea surface temperature (>21°C) and increased primary productivity. Our results indicate strong seasonal and interannual philopatry to the waters of Virginia (summer foraging habitat) and North Carolina (winter habitat). We suggest that the waters of Virginia and North Carolina provide important seasonal habitat and serve as a seasonal migratory pathway for immature loggerhead sea turtles. North Carolina’s Cape Hatteras acts as a seasonal “migratory bottleneck” for this species; special management consideration should be given to this region. Six turtles spent time farther from the continental shelf. Three entered the Gulf Stream near Cape Hatteras, traveling in the current to the northwest Atlantic. Two of these turtles remained within an oceanic habitat from 1 to 3 years and were associated with mesoscale features and frontal systems. The ability of large benthic subadults to resume an oceanic lifestyle for extended periods indicates plasticity in habitat use and migratory strategies. Therefore, traditional life history models for loggerhead sea turtles should be reevaluated.     

Foraging areas differentially affect reproductive output and interpretation of trends in abundance of loggerhead turtles

Diet items and habitat constitute some of the environmental resources that may be used differently by individuals within a population. Long-term fidelity by individuals to particular resources exemplifies individual specialization, a phenomenon that is becoming increasingly recognized across a wide range of species. Less is understood about the consequences of such specialization. Here, we investigate the effects of differential foraging ground use on reproductive output in 183 loggerhead sea turtles (Caretta caretta) nesting at Wassaw Island, Georgia (31.89°N, 80.97°W), between 2004 and 2011 with resulting possible fitness effects. Stable isotope analysis was used to assign the adult female loggerheads to one of three foraging areas in the Northwest Atlantic Ocean. Our data indicate that foraging area preference influences the size, fecundity, and breeding periodicity of adult female loggerhead turtles. We also found that the proportion of turtles originating from each foraging area varied significantly among the years examined. The change in the number of nesting females across the years of the study was not a result of uniform change from all foraging areas. We develop a novel approach to assess differential contributions of various foraging aggregations to changes in abundance of a sea turtle nesting aggregation using stable isotopes. Our approach can provide an improved understanding of the influences on the causes of increasing or decreasing population trends and allow more effective monitoring for these threatened species and other highly migratory species.     

A model of area fidelity,nomadism, and distribution patterns of loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) in the Mediterranean Sea

Sea turtle tagging carried out in Italy in the period 1981–2006 resulted in 125 re-encounters of loggerhead turtles (Caretta caretta) after a mean of 2.5 years, from different marine areas in the Mediterranean. At first finding, turtles ranged 25–83 cm of curved carapace length. Data were analyzed according to size, area, habitat type, season, in order to provide indication of movement patterns. When integrated with other information, results indicate that: (1) a part of turtles in the oceanic stage show a nomad behavior with movements among different oceanic areas; (2) another part show fidelity to an oceanic area; (3) turtles in the neritic stage show fidelity to neritic areas, and once settled to one area, change to other neritic areas is unlikely; (4) nomad oceanic turtles are significantly larger than sedentary ones, and also larger than turtles found in neritic areas; it is hypothesized that these could be Atlantic turtles that eventually leave the Mediterranean; (5) ecological transition from oceanic to neritic habitats occurs at a wide range of sizes, and some turtles may have a very brief oceanic stage; (6) turtles in the oceanic stage are more likely to recruit to neritic areas close to their oceanic areas than to distant ones; (7) part of turtles from some Mediterranean nesting beaches might frequent a relatively limited area range, including both oceanic and neritic areas; (8) in most of the Mediterranean, latitudinal seasonal migrations are unlikely. A general model of movement patterns of loggerhead turtles in the Mediterranean is proposed.     

Foraging habitats and migration corridors utilized by a recovering subpopulation of adult female loggerhead sea turtles: implications for conservation

From 1998 to 2008, 68 adult female loggerhead sea turtles (Caretta caretta) were instrumented with platform transmitter terminals at nesting beaches in Georgia, North Carolina (NC) and South Carolina (SC) on the East Coast of the United States of America (30°48′N, 81°28′W to 33°51′N, 77°59′W). The majority of post-nesting loggerheads (N = 42, 62 %) migrated to foraging habitats in the Mid-Atlantic Bight during May–October, with a subsequent migration occurring during November–March to foraging habitats south of Cape Hatteras, NC. Nine (13 %) loggerheads initially foraged in the near-shore, coastal areas of the South Atlantic Bight, but moved to offshore habitats—closer to the Gulf Stream—during November–March, while fourteen (21 %) loggerheads remained in foraging areas along the mid-continental shelf off of the eastern coast of Florida and/or continued southward to Florida Bay and the Bahamas. The present study delineates important, post-nesting foraging habitats and migration corridors where loggerheads may interact with commercial fisheries—providing managers opportunities to develop and implement optimally effective conservation actions for the recovery of this threatened species.     

Post-nesting migrations of loggerhead sea turtles in the Gulf of Mexico: dispersal in highly dynamic conditions

This study is the first report of post-nesting migrations of loggerhead sea turtles (Caretta caretta) nesting in Sarasota County (Florida, USA), their most important rookery in the Gulf of Mexico (GOM). In total, 28 females (curved carapace length CCL between 82.2 and 112.0 cm) were satellite-tracked between May 2005 and December 2007. Post-nesting migrations were completed in 3–68 days (mean ± SD = 23 ± 16 days). Five different migration patterns were observed: six turtles remained in the vicinity of their nesting site while the other individuals moved either to the south-western part of the Florida Shelf (n = 9 turtles), the Northeast GOM (n = 2 turtles), the South GOM (Yucatán Shelf and Campeche Bay, Mexico, and Cuba; n = 5 turtles) or the Bahamas (n = 6 turtles). In average, turtles moved along rather straight routes over the continental shelf but showed more indirect paths in oceanic waters. Path analyses coupled with remote sensing oceanographic data suggest that most of long-distance migrants reached their intended foraging destinations but did not compensate for the deflecting action of ocean currents. While six out of seven small individuals (CCL < 90 cm) remained on the Florida Shelf, larger individuals showed various migration strategies, staying on the Florida Shelf or moving to long-distance foraging grounds. This study highlights the primary importance the Western Florida Shelf in the management of the Florida Nesting Subpopulation, as well as the need of multi-national effort to promote the conservation of the loggerhead turtle in the Western Atlantic. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Habitat use by loggerhead sea turtles <Emphasis Type="Italic">Caretta caretta</Emphasis> off the coast of eastern Spain results in a high vulnerability to neritic fishing gear

Previous studies of loggerhead sea turtles have concluded that drifting longlines were the main threat for immature specimens in the western Mediterranean, because immature loggerhead sea turtles mainly inhabit oceanic waters. However, recent aerial surveys have revealed large numbers of immature loggerhead sea turtles over the continental shelf of eastern mainland Spain, where turtles are exposed to neritic fishing gears but not to drifting longlines. We satellite-tracked seven loggerhead sea turtles (minimum straight carapace length (SCLmin) range: 36.5–55.0 cm) to assess whether the turtles in this region are vagrants from the adjoining oceanic regions or whether these loggerheads mostly inhabit the continental shelf. Satellite-tracking revealed that six of the tagged turtles avoided the oceanic realm and made extended use of the continental shelf, whereas only one individual could be considered a true vagrant as it avoided the continental shelf and primarily used the oceanic habitat. These results are in sharp contrast with those previously reported for immature loggerhead sea turtles of similar size from the south-western Mediterranean and fit well a relaxed ontogenic model that was recently proposed for loggerhead sea turtles in the central Mediterranean. Furthermore, these results demonstrate the vulnerability of loggerhead sea turtles of eastern mainland Spain to neritic fishing gears, as three of the seven turtles died and one was bycaught incidentally while being tracked over the continental shelf.     

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.     

Regional and annual variation in plasma steroids and metabolic indicators in female green turtles, <Emphasis Type="Italic">Chelonia mydas</Emphasis>

Variation in environmental conditions at a foraging area or at a nesting rookery has the potential to impact reproductive output of green sea turtles (Chelonia mydas) by affecting food resources or the nesting substrate. In this paper we test whether turtles‘ physiological characteristics reflect variation in relevant environmental conditions. We did this by profiling metabolic and hormonal markers among (1) non-vitellogenic females from three different foraging areas and (2) nesting females from different rookeries and breeding seasons. Among the non-vitellogenic females, the highest plasma triglyceride concentrations (4.29 mmol/l) and the lowest plasma cholesterol concentrations (1.27 mmol/l) were found in non-vitellogenic females residing in Moreton Bay during the El Niño year of 1997. Furthermore, during 1997, these Moreton Bay females had higher plasma triglyceride and lower cholesterol concentrations than those recorded in non-vitellogenic females foraging at Heron Reef (triglyceride 1.22 mmol/l and cholesterol 4.53 mmol/l) and Shoalwater Bay (triglyceride 1.69 mmol/l and cholesterol 3.50 mmol/l) in the same year. Among nesting turtles, those nesting at Raine Island had low mean plasma triglyceride concentrations during the high density 1996 nesting season. For those nesting at Heron Island, the mean triglyceride concentrations were the lowest during the 1997 nesting season. This is the first time that hormone and metabolic markers have been used in concert to compare the physiological condition of nesting and foraging sea turtles and its relationship with the environment. Collectively, our data indicate that variation in the environmental conditions at both foraging and nesting areas are reflected at a physiological level. Moreover, our study indicates that turtles feeding during El Niño years are able to attain higher levels of body condition, and that physiological data combined with morphometric data is a useful proxy for assessing the condition of turtles in foraging areas.     

Satellite tracking of migrating loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) displaced in the open sea

Loggerhead turtles (Caretta caretta) are known to migrate towards fixed, individually-specific residential feeding grounds. To study their spatial behaviour and their navigational ability, five loggerheads nesting in South Africa were captured when about to start their postnesting migration and tracked by satellite after having been displaced from their usual migratory route. The first turtle, released south of Madagascar about 1,148 km from the capture site, moved west up to mainland Africa and then reached her feeding grounds by following the coast. A second turtle, released farther away (2,140 km) close to La Réunion Island, stopped for some time on the Madagascar east coast, then turned southwards to round the island and regain the African mainland in the northwest, without however allowing us to establish the location of her residential grounds. Three other turtles were released off the Tanzanian coast, 2,193 km north of their nesting area, at the northern edge of the distribution of the feeding grounds along the African coast. All of them headed north, and one turtle found her residential grounds located north of the release site. The other two females started long-distance oceanic wanderings in which they crossed nearly the entire Indian Ocean, apparently being transported by the sea currents of the region. We conclude that adult loggerhead turtles are apparently unable to compensate for the displacement and can return to a pelagic life style characteristic of juvenile turtles. These findings suggest that South African loggerheads rely on simple orientation mechanisms, such as the use of the coastline, as a guide, and compass orientation, possibly integrated by spatiotemporal programmes and/or acquired maps of familiar sites.Communicated by R. Cattaneo-Vietti, Genova     

Are anthropogenic factors affecting nesting habitat of sea turtles? The case of Kanzul beach,Riviera Maya-Tulum (Mexico)

Marine coast modification and human pressure affects many species, including sea turtles. In order to study nine anthropogenic impacts that might affect nesting selection of females, incubation and hatching survival of loggerhead (Caretta caretta) and green turtle (Chelonia mydas), building structures were identified along a 5.2 km beach in Kanzul (Mexico). A high number of hotels and houses (88; 818 rooms), with an average density of 16.6 buildings per kilometer were found. These buildings form a barrier which prevents reaching the beach from inland, resulting in habitat fragmentation. Main pressures were detected during nesting selection (14.19% of turtle nesting attempts interrupted), and low impact were found during incubation (0.77%) and hatching (4.7%). There were three impacts defined as high: beach furniture that blocks out the movement of hatchlings or females, direct pressure by tourists, and artificial beachfront lighting that can potentially mislead hatchlings or females. High impacted areas showed lowest values in nesting selection and hatching success. Based on our results, we suggest management strategies to need to be implemented to reduce human pressure and to avoid nesting habitat loss of loggerhead and green turtle in Kanzul, Mexico.     

Migration,distribution, and diving behavior of adult male loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) following dispersal from a major breeding aggregation in the Western North Atlantic

Sixteen satellite-tagged adult male loggerhead sea turtles (Caretta caretta) dispersed widely from an aggregation near Port Canaveral, Florida, USA (28°23′N, −80°32′W) after breeding. Northbound males migrated further (990 ± 303 km) than southbound males (577 ± 168 km) and transited more rapidly (median initial dive duration = 6 (IQR = 4–16) versus 19 (IQR = 10–31) min, respectively).. Migration occurred along a depth corridor (20–40 m) except where constricted by a narrow continental shelf width. Males foraged in areas 27 ± 41 km² day⁻¹ at locations <1–80 km from shore for 100.1 ± 60.6 days, with variability in foraging patterns not explained by turtle size or geography. Post-breeding dispersal patterns were similar to patterns reported for adult female loggerhead sea turtles in this region and adult male loggerhead sea turtles elsewhere in the northern hemisphere; however, foraging ground distributions were most similar to adult female loggerhead sea turtles in this region.