A review of long-distance movements by marine turtles, and the possible role of ocean currents

Sea turtle movements often occur in open-sea unsheltered areas, and are therefore likely to be influenced by major oceanographic processes. Only recently has work started to examine the possible relationships of these movements with dynamic oceanic features, and consequently a clear picture of such interaction is only available in a few cases. Newborn sea turtles are thought to rely on oceanic currents to reach their pelagic nursery habitats. The actual extent and timing of these developmental migrations are known for only a few populations, but these movements probably last several years and range over thousands of km. Large juveniles that have been tracked during their pelagic stage were found to make long-distance movements, sometimes swimming against the prevailing currents. Older juveniles of most species leave the pelagic habitat to recruit to neritic developmental habitats. This is a very poorly documented phase of the sea turtle life-cycle, and the few available indications show that turtles may have to swim actively for enormous distances to counterbalance their previous drift with the current. The course and extent of adult postnesting migrations vary greatly among different turtle species, but two main patterns are evident. Some species, like green, hawksbill and loggerhead turtles, shuttle between the nesting beach and a specific feeding area used for the entire inter-reproductive period. In these cases, individuals swim, rather than drift, to complete their journeys, with possible advection due to currents sometimes helping them to quickly reach their target, but sometimes providing navigational challenges. Other species such as the olive ridley and the leatherback turtle, leave the coastal nesting areas to reach the pelagic environment where they forage, and perform wandering movements. Major oceanographic processes (such as main currents and eddies) have been recently shown to have a remarkable influence on leatherback movements, making it questionable whether these journeys are to be considered migrations or, rather, prolonged stays in vast feeding areas.

Habitat utilization by juvenile hawksbill turtles (Eretmochelys imbricata, Linnaeus, 1766) around a shallow water coral reef

Contemporary studies of sea turtle diving behaviour are generally based upon sophisticated techniques such as the attachment of time depth recorders. However, if the risks of misinterpretation are to be minimized, it is essential that electronic data are analysed in the light of first-hand observations. To this aim, we set out to make observations of juvenile hawksbill turtles (Eretmochelys imbricata, Linnaeus, 1766) foraging and resting in a shallow water coral reef habitat around the granitic Seychelles (4°'S, 55°'E). Data were collected from six study sites characterized by a shallow reef plateau (<5 m) and a flat sandy area at the base of the reef face (<10 m). Observation data were categorized into the following behaviours: (1) stationary foraging; (2) active foraging; (3) resting; and (4) assisted resting. Central to this investigation was the development of a technique for accurately estimating the size of sea turtles in situ based upon previously tested techniques for reef fishes. This revealed that through calibration, the curved carapace length (CCL) of marine turtles can be consistently estimated to within 10 cm of their actual size. Although rudimentary, this has advantages for assessing the residency or absence of specific life history stages from particular environments. Indeed, our data supported previous claims that following the reproductive season, adult hawksbills in the region may move away from the nesting beaches to alternative foraging grounds whilst immature turtles (following the pelagic juvenile stage) may opt to reside in areas close to their natal beaches. With regards to habitat utilization, juvenile hawksbills displayed an alternating pattern of short, shallow foraging dives followed by deeper, longer resting dives. These findings are consistent with previous electronic studies of free-range diving in this species and suggest that the maximization of resting duration may be an important factor driving this behaviour.

Movement patterns of green turtles in Brazilian coastal waters described by satellite tracking and flipper tagging

The movements of 8 green turtles Chelonia mydas in Brazilian coastal waters were tracked using transmitters linked to the Argos system for periods of between 1 and 197 d. These were the first tracking data gathered on juveniles of this species in this important foraging ground. Information was integrated with that collected over a decade using traditional flipper-tagging methods at the same site. Both satellite telemetry and flipper tagging suggested that turtles undertook 1 of 3 general patterns of behaviour: pronounced long range movements (>100 km), moderate range movements (<100 km) or extended residence very close to the capture/release site. There seemed to be a general tendency for the turtles recaptured/tracked further afield to have been among the larger turtles captured. Satellite tracking of 5 turtles which moved from the release site showed that they moved through coastal waters; a factor which is likely to predispose migrating turtles to incidental capture as a result of the prevailing fishing methods in the region. The movements of the 3 turtles who travelled less than 100 km from the release site challenge previous ideas relating to home range in green turtles feeding in sea grass pastures. We hypothesise that there may be a fundamental difference in the pattern of habitat utilisation by larger green turtles depending on whether they are feeding on seagrass or macroalgae. Extended tracking of 2 small turtles which stayed near the release point showed that small juvenile turtles, whilst in residence in a particular feeding ground, can also exhibit high levels of site-fidelity with home ranges of the order of several square kilometers.

Change in body mass associated with long-term fasting in a marine reptile : the case of green turtles (Chelonia mydas) at Ascension Island

Female green sea turtles (Chelonia mydas) nesting at Ascension Island (7°57'S, 14°22'W) in the middle of the Atlantic Ocean had a mean body mass (post oviposition) of 166.3 kg (range 107.5–243.5 kg, n = 119). Individuals lost mass slowly during the nesting season (mean mass loss 0.22 kg·d–1, n = 14 individuals weighed more than once). Gut-content analysis and behavioural observations indicated a lack of feeding. Females of equivalent-sized pinniped species that also do not feed while reproducing (nursing pups) on islands lose mass about 17 times faster. This comparatively low rate of mass loss by green turtles probably reflects their ectothermic nature and, consequently, their low metabolic rate. We estimate that a female turtle would lose only 19% of her body mass during the 143-day, 4400-km round trip from Brazil if she did not eat, laid 3 clutches of eggs, and lost 0.22 kg·d–.

Long-term satellite telemetry of the movements and habitat utilisation by green turtles in the Mediterranean

There is a relative paucity of data regarding the at-sea distribution and behaviour of marine turtles. This is especially true for the critically endangered green turtle Chelonia mydas population in the Mediterranean. Six adult female green turtles were equipped with satellite transmitters and tracked for periods of between 28 and 293 d following their final nesting of the season in northern Cyprus. Data elucidated hitherto unknown migratory pathways and highlighted the importance of North African coastal waters as feeding habitat for adults of this species. For three individuals, instruments transmitted detailed information on dive depth, dive duration and water temperature which afforded novel insights into behaviour during different stages of migration, feeding in the foraging grounds and most remarkably, during a period of midwinter diapause when water temperatures were generally below 25°C. Turtles showed fidelity to specific shallow inshore feeding areas and moved offshore to deeper wintering sites.

Temperature-dependent sex determination of Ascension Island green turtles

We examined the role played by temperature in the duration of incubation and sex ratio of green turtle hatchlings at Ascension Island, one of the most important green turtle rookeries in the Atlantic. Temperature at control sites at nest depth and in 39 green turtle nests was measured using small temperature recording devices. The sex ratio of hatchlings was ascertained in a sub-sample of monitored nests allowing the description of the relationship between intranest temperature and hatchling sex ratio, demonstrating a pivotal incubation temperature of 28.8°C. The seasonal profile in sex ratio of hatchlings produced on all nesting beaches at Ascension Island was estimated, showing that a female-biased sex ratio would be expected with a female:male ratio of the order of 3:1. The use of nest temperature, air temperature, sand temperature at control sites, and incubation duration as proxies to estimate hatchling sex ratio are discussed.

The diving behaviour of green turtles undertaking oceanic migration to and from Ascension Island: dive durations, dive profiles and depth distribution

Satellite telemetry was used to record the submergence duration of green turtles (Chelonia mydas) as they migrated from Ascension Island to Brazil (N=12 individuals) while time/depth recorders (TDRs) were used to examine the depth distribution and dive profiles of individuals returning to Ascension Island to nest after experimental displacement (N=5 individuals). Satellite telemetry revealed that most submergences were short (<5 min) but that some submergences were longer (>20 min), particularly at night. TDRs revealed that much of the time was spent conducting short (2–4 min), shallow (approximately 0.9–1.5 m) dives, consistent with predictions for optimisation of near-surface travelling, while long (typically 20–30 min), deep (typically 10–20 m) dives had a distinctive profile found in other marine reptiles. These results suggest that green turtles crossing the Atlantic do not behave invariantly, but instead alternate between periods of travelling just beneath the surface and diving deeper. These deep dives may have evolved to reduce silhouetting against the surface, which would make turtles more susceptible to visual predators such as large sharks.

Changes in behaviour during the inter-nesting period and post-nesting migration for Ascension Island green turtles

Satellite transmitters were attached to green turtles Chelonia mydas while they were nesting on Ascension Island in the South Atlantic (7°57'S, 14°22'W) and individuals were subsequently monitored during the inter-nesting period and the post-nesting migration to Brazil. During the inter-nesting period, data from the transmitters suggested that turtles generally stayed within 5 km of the nesting beach on which they had originally been observed. During both the inter-nesting period and migration, turtles were submerged the vast majority (>95%) of the time, suggesting that they neither basked at the surface nor drifted passively during migration to any great extent. There was a clear dichotomy in submergence behaviour, with submergences tending to be of short duration during post-nesting migration (mean = 7.3 min, 3318 h of data from 5 individuals) and of longer duration during the inter-nesting period (mean = 22.1 min, 714 h of data from 5 different individuals). As submergence duration is generally linked to activity levels in sea turtles, this pattern suggests that turtles were comparatively inactive during the inter-nesting period and comparatively active during migration. During both the inter-nesting period and the post-nesting migration, diel submergence patterns were detected with dive duration tending to be longer at night. As the turtles migrated WSW from Ascension Island, there was a reduction in their speed of travel. A numerical model of the near-surface currents suggested that this reduction was associated with the weakening of the WSW flow of the prevailing South Atlantic Equatorial Current.

Long-term thermal conditions on the nesting beaches of green turtles on Ascension Island

On 2 of the major nesting beaches used by green turtles Chelonia mydas on Ascension Island, we measured the sand temperature at nest depths throughout the year. For both beaches, the sand temperature was strongly correlated (r2 >= 0.94) with air temperature. We therefore used past records of air temperature to reconstruct sand temperatures on the different beaches throughout the nesting season between 1985 and 1998. This analysis showed that inter-annual differences in sand temperature were small and, while there were consistent thermal changes during the nesting season, over the 14 yr there was little overlap in the temperatures on the 2 beaches, with one being 2.6°C warmer, on average, than the other. This work suggests that inter-beach thermal variation is the major mechanism by which a range of incubation temperatures are realised on Ascension Island and hence is likely to facilitate the production of hatchlings of both sexes.

The navigational feats of green sea turtles migrating from Ascension Island investigated by satellite telemetry

Previous tagging studies of the movements of green turtles (Chelonia mydas) nesting at Ascension Island have shown that they shuttle between this remote target in the Atlantic Ocean and their feeding grounds on the Brazilian coast, a distance of 2300 km or more. Since a knowledge of sea turtle migration routes might allow inferences on the still unknown navigational mechanisms of marine animals, we tracked the postnesting migration of six green turtle females from Ascension Island to Brazil. Five of them reached the proximity of the easternmost stretch of the Brazilian coast, covering 1777 to 2342 km in 33 to 47 days. Their courses were impressively similar for the first 1000 km, with three turtles tracked over different dates following indistinguishable paths for the first 300 km. Only the sixth turtle made some relatively short trips in different directions around Ascension. The tracks show that turtles (i) are able to maintain straight courses over long distances in the open sea; (ii) may perform exploratory movements in different directions; (iii) appropriately correct their course during the journey according to external information; and (iv) initially keep the same direction as the west–south–westerly flowing current, possibly guided by chemical cues.

Unravelling migratory connectivity in marine turtles using multiple methods

1. Comprehensive knowledge of the fundamental spatial ecology of marine species is critical to allow the identification of key habitats and the likely sources of anthropogenic threats, thus informing effective conservation strategies. 2. Research on migratory marine vertebrates has lagged behind many similar terrestrial animal groups, but studies using electronic tagging systems and molecular techniques offer great insights. 3. Marine turtles have complex life history patterns, spanning wide spatio-temporal scales. As a result of this multidimensional complexity, and despite extensive effort, there are no populations for which a truly holistic understanding of the spatial aspects of the life history has been attained. There is a particular lack of information regarding the distribution and habitats utilized during the first few years of life. 4. We used satellite tracking technology to track individual turtles following nesting at the green turtle Chelonia mydas nesting colony at Poilão Island, Guinea Bissau; the largest breeding aggregation in the eastern Atlantic. 5. We further contextualize these data with pan-Atlantic molecular data and oceanographic current modelling to gain insights into likely dispersal patterns of hatchlings and small pelagic juveniles. 6. All adult turtles remained in the waters of West Africa, with strong connectivity demonstrated with Banc D’Arguin, Mauritania. 7. Despite shortcomings in current molecular markers, we demonstrate evidence for profound sub-structuring of marine turtle stocks across the Atlantic; with a high likelihood based on oceanographic modelling that most turtles from Guinea-Bissau are found in the eastern Atlantic. 8. Synthesis and applications. There is an increased need for a better understanding of spatial distribution of marine vertebrates demonstrating life histories with spatio-temporal complexity. We propose the synergistic use of the technologies and modelling used here as a working framework for the future rapid elucidation of the range and likely key habitats used by the different life stages from such species.

Global research priorities for sea turtles : informing management and conservation in the 21st century

Over the past 3 decades, the status of sea turtles and the need for their protection to aid population recovery have increasingly captured the interest of government agencies, non-governmental organisations (NGOs) and the general public worldwide. This interest has been matched by increased research attention, focusing on a wide variety of topics relating to sea turtle biology and ecology, together with the interrelations of sea turtles with the physical and natural environments. Although sea turtles have been better studied than most other marine fauna, management actions and their evaluation are often hindered by the lack of data on turtle biology, human–turtle interactions, turtle population status and threats. In an effort to inform effective sea turtle conservation a list of priority research questions was assembled based on the opinions of 35 sea turtle researchers from 13 nations working in fields related to turtle biology and/or conservation. The combined experience of the contributing researchers spanned the globe as well as many relevant disciplines involved in conservation research. An initial list of more than 200 questions gathered from respondents was condensed into 20 metaquestions and classified under 5 categories: reproductive biology, biogeography, population ecology, threats and conservation strategies.

Leatherback turtles satellite-tagged in European waters

The North Atlantic is considered a stronghold for the critically endangered leatherback sea turtle. However, limited information exists regarding the movements of individuals to and from the seas off Europe’s northwesterly fringe, an area where leatherbacks have been historically sighted for the past 200 yr. Here, we used satellite telemetry to record the movements and behaviour of 2 individuals bycaught in fisheries off the southwest coast of Ireland. The turtle T1 (tagged 1 September 2005; female; tracked 375 d) immediately travelled south via Madeira and the Canaries, before residing in West African waters for 3 mo. In spring, T1 migrated north towards Newfoundland where transmissions ceased. T2 (29 June 2006; male; 233 d) travelled south for a short period before spending 66 d west of the Bay of Biscay, an area previously asserted as a high-use area for leatherbacks. This prolonged high latitude summer residence corresponded with a mesoscale feature evident from satellite imagery, with the implication that this turtle had found a rich feeding site. A marked change in dive behaviour was apparent as the turtle exited this feature and provided useful insights on leatherback diving behaviour. T2 headed south in October 2006, and performed the deepest-ever dive recorded by a reptile (1280 m) southwest of Cape Verde. Unlike T1, T2 swam southwest towards Brazil before approaching the major nesting beaches of French Guiana and Surinam. Importantly, these tracks document the movement of leatherbacks from one of the remotest foraging grounds in the North Atlantic.

The role of infrequent and extraordinary deep dives in leatherback turtles (Dermochelys coriacea)

Infrequent and exceptional behaviours can provide insight into the ecology and physiology of a particular species. Here we examined extraordinarily deep (300–1250 m) and protracted (>1h) dives made by critically endangered leatherback turtles (Dermochelys coriacea) in the context of three previously suggested hypotheses: predator evasion, thermoregulation and exploration for gelatinous prey. Data were obtained via satellite relay data loggers attached to adult turtles at nesting beaches (N=11) and temperate foraging grounds (N=2), constituting a combined tracking period of 9.6 years (N=26,146 dives) and spanning the entire North Atlantic Ocean. Of the dives, 99.6% (N=26,051) were to depths <300 m with only 0.4% (N=95) extending to greater depths (subsequently termed `deep dives'). Analysis suggested that deep dives: (1) were normally distributed around midday; (2) may exceed the inferred aerobic dive limit for the species; (3) displayed slow vertical descent rates and protracted durations; (4) were much deeper than the thermocline; and (5) occurred predominantly during transit, yet ceased once seasonal residence on foraging grounds began. These findings support the hypothesis that deep dives are periodically employed to survey the water column for diurnally descending gelatinous prey. If a suitable patch is encountered then the turtle may cease transit and remain within that area, waiting for prey to approach the surface at night. If unsuccessful, then migration may continue until a more suitable site is encountered. Additional studies using a meta-analytical approach are nonetheless recommended to further resolve this matter.