Fidelity to foraging sites, consistency of migration routes and habitat modulation of home range by sea turtles

Aim  Resources can shape patterns of habitat utilization. Recently a broad foraging dichotomy between oceanic and coastal sites has been revealed for loggerhead sea turtles (Caretta caretta). Since oceanic and coastal foraging sites differ in prey availability, we might expect a gross difference in home-range size across these habitats. We tested this hypothesis by equipping nine adult male loggerhead sea turtles with GPS tracking devices. Location  National Marine Park of Zakynthos (NMPZ) Greece, central and eastern Mediterranean (Adriatic, Ionian and Aegean seas). Methods  In 2007, 2008 and 2009, Fastloc GPS-Argos transmitters were attached to nine male loggerheads. In addition, a Sirtrack PTT unit was attached to one male in 2007. Four of the turtles were tracked on successive years. We filtered the GPS data to ensure comparable data volumes. Route consistency between breeding and foraging sites of the four re-tracked turtles was conducted. Foraging site home range areas and within site movement patterns were investigated by the fixed kernel density method. Results  Foraging home range size ranged between circa 10 km2 at neritic habitats (coastal and open-sea on the continental shelf) to circa 1000 km2 at oceanic sites (using 90% kernel estimates), the latter most probably reflecting sparsely distributed oceanic prey. Across different years individuals did not follow exactly the same migration routes, but did show fidelity to their previous foraging sites, whether oceanic or neritic, with accurate homing in the final stages of migration. Main conclusions  The broad distribution and diverse life-history strategies of this population could complicate the identification of priority marine protected areas beyond the core breeding site.

Microhabitat selection by sea turtles in a dynamic thermal marine environment

1Reproductive fitness is often compromised at the margins of a species’ range due to sub-optimal conditions.2Set against this backdrop, the Mediterranean's largest loggerhead sea turtle (Caretta caretta) rookery at Zakynthos (Greece) presents a conundrum, being at a very high latitude for this species, yet hosting a high concentration of nesting.3We used visual surveys combined with global positioning system (GPS) tracking to show that at the start of the breeding season, individuals showed microhabitat selection, with females residing in transient patches of warm water. As the sea warmed in the summer, this selection was no longer evident.4As loggerhead turtles are ectothermic, this early season warm-water selection presumably speeds up egg maturation rates before oviposition, thereby allowing more clutches to be incubated when sand conditions are optimal during the summer.5Active selection of warm waters may allow turtles to initiate nesting at an earlier date.

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.

Influence of ocean currents on long-distance movement of leatherback sea turtles in the Southwest Indian Ocean

Leatherback turtles Dermochelys coriacea spend most of their life in oceanic environments, whose physical and biological characteristics are primarily forged by sea current circulation. Water mass movements can mechanically act on swimming turtles, thus determining their routes, and can differentially distribute their planktonic prey. By integrating satellite tracking data with contemporaneous remote-sensing information, we analysed the post-nesting journeys of 9 leatherbacks with respect to oceanographic surface conditions. Tracked turtles showed large variations in migration routes and in final destinations, apparently without heading for specific foraging areas. Their complex tracks spread over wide regions around South Africa. Leatherbacks were greatly influenced by the currents encountered during their movements, with their trajectories displaying curves or revolutions in the presence of (and in accordance with) rotating water masses. An impressive similarity was observed between large parts of the turtle routes and those of surface drifters tracked in the same regions. Finally, leatherbacks remained associated for long periods with specific oceanographic features, which most probably offered them profitable foraging opportunities. These results agree with previous findings in showing a strong influence of oceanic currents and mesoscale features on the movements of South African leatherbacks, and additionally identify the role of current-related features in causing the observed route variability and in determining high-quality foraging hotspots for leatherbacks moving in the ocean.

Behavioral inference of diving metabolic rate in free-ranging leatherback turtles

Good estimates of metabolic rate in free‐ranging animals are essential for understanding behavior, distribution, and abundance. For the critically endangered leatherback turtle (Dermochelys coriacea), one of the world’s largest reptiles, there has been a long‐standing debate over whether this species demonstrates any metabolic endothermy. In short, do leatherbacks have a purely ectothermic reptilian metabolic rate or one that is elevated as a result of regional endothermy? Recent measurements have provided the first estimates of field metabolic rate (FMR) in leatherback turtles using doubly labeled water; however, the technique is prohibitively expensive and logistically difficult and produces estimates that are highly variable across individuals in this species. We therefore examined dive duration and depth data collected for nine free‐swimming leatherback turtles over long periods (up to 431 d) to infer aerobic dive limits (ADLs) based on the asymptotic increase in maximum dive duration with depth. From this index of ADL and the known mass‐specific oxygen storage capacity (To2) of leatherbacks, we inferred diving metabolic rate (DMR) as . We predicted that if leatherbacks conform to the purely ectothermic reptilian model of oxygen consumption, these inferred estimates of DMR should fall between predicted and measured values of reptilian resting and field metabolic rates, as well as being substantially lower than the FMR predicted for an endotherm of equivalent mass. Indeed, our behaviorally derived DMR estimates ( mL O2 min−1 kg−1) were times the resting metabolic rate measured in unrestrained leatherbacks and times the average FMR for a reptile of equivalent mass. These DMRs were also nearly one order of magnitude lower than the FMR predicted for an endotherm of equivalent mass. Thus, our findings lend support to the notion that diving leatherback turtles are indeed ectothermic and do not demonstrate elevated metabolic rates that might be expected due to regional endothermy. Their capacity to have a warm body core even in cold water therefore seems to derive from their large size, heat exchangers, thermal inertia, and insulating fat layers and not from an elevated metabolic rate.

Meta-analysis of movements in Atlantic leatherback turtles during the nesting season : conservation implications

Despite decades of conservation efforts on the nesting beaches, the critical status of leatherback turtles shows that their survival predominantly depends on our ability to reduce at-sea mortality. Although areas where leatherbacks meet fisheries have been identified during the long distance movements between 2 consecutive nesting seasons, hot-spots of lethal interactions are still poorly defined within the nesting season, when individuals concentrate close to land. Here we report movements of satellite-tracked gravid leatherback turtles during the nesting season in Western Central Africa, South America and the Caribbean Sea, which account for about 70% of the world population. We show that during and at the end of the nesting season, leatherback turtles have the propensity to remain over the continental shelf, yet sometimes perform extended movements and may even nest in neighbouring countries. Leatherbacks exploit coastal commercial fishing grounds and face substantial accidental capture by regional coastal fisheries (e.g. at least 10% in French Guiana). This emphasises the need for regional conservation strategies to be developed at the ocean scale—both at sea and on land—to ensure the survival of the last leatherback turtles.

Flipper beat frequency and amplitude changes in diving green turtles, Chelonia mydas

Using a turtle-borne camera system, changing flipper beat frequency and amplitude were measured in five diving green turtles (Chelonia mydas Linnaeus 1758) in the Bahía de los Angeles, Mexico (28°58′N, 113°33′W). These observations were made between June and August 2002. Turtles worked hardest (i.e., had the highest flipper beat frequency and amplitude) at the start of descents when positive buoyancy is predicted to oppose their forward motion. During the later part of descents, turtles worked less hard in line with opposing buoyancy forces being reduced. For example, flipper beat frequency declined from about 60–80 beats min−1 at the start of descent to around 25–40 beats min−1 after 30 s of the descent. At the start of ascents the flipper beat frequency was around 30 beats min−1, lower than on descent, and declined as the ascent progressed with often passive gliding for the final few meters to the surface. This pattern of effort during diving appears to apply across a range of marine reptiles, birds and mammals suggesting that graded effort during descent and ascent is an optimum solution to minimising the cost of transport during diving.

Overwintering behaviour in sea turtles : dormancy is optional

Thirteen loggerhead turtles Caretta caretta were released (10 from Naples, Italy, 2 from Monastir, Tunisia, 1 from Gallipoli, South Italy) with satellite relay data loggers (SRDL) to elucidate their overwintering behaviour. Nine turtles were successfully tracked throughout the winter, while 4 SRDLs failed to transmit after short deployment periods. Of these 9, 4 remained within 80 km of the release site, 3 travelled to a distant overwintering site, and 2 continued to move and did not remain within 80 km of a specific site. Apart from these differences, all turtles stayed near the coast and dedicated most of their time to dives lasting 3 h and longer. Maximum dive durations ranged from 270 to 480 min and were highly correlated with water temperatures, which fell below the supposed 15°C threshold for sea turtle hibernation in all overwintering sites. Median dive depths were between 4 and 24 m and were, thus, well within the mixed layer, as revealed by temperature profiles, which also were relayed by the SRDLs. No evidence was found that the turtles preferred warmer temperatures to overwinter in, because the range of temperature was very narrow on both the horizontal and the vertical scale of their movements. Despite the long resting phases and the low temperatures (minimum = 11.8°C) all turtles retained activity to some degree, at least to commute between the depth of resting and the surface to breathe. While the degree of winter dormancy is certainly affected by temperature, turtles were by no means obligatory hibernators, and their ability to move and even forage during the winter may be important for their growth and maturation rates, as well as their reproductive output.

Female–female aggression : structure of interaction and outcome in loggerhead sea turtles

Aggressive behaviour between females of the same species is not widely documented, particularly in marine vertebrates. During a 3 yr in-water survey at the temperate loggerhead sea turtle Caretta caretta breeding area of Zakynthos, Greece, female–female interactions comprised 4% of all female loggerhead sighting events (n = 60 out of 1449 events). Male–female interactions comprised an additional 4% of sighting events, while 92% were of solitary females. The structure of interactions was analysed for 58 of these sighting events, each lasting an average of 3.4 min (SD ± 1) and comprising a total of 3.1 h observation time. We found that interactions involved ritualized escalation in behaviour from passive threat displays (e.g. head–tail circling) to aggressive combat (e.g. sparring). We suggest that circling individuals evaluate opponent size, sparring individuals test opponent strength, and that the positioning of the prehensile tail signals motivational intent to either escalate or abort. The presence of intruder females triggered a passive response in 100% of events involving basking and swimming turtles (n = 19); although residents resting on the seabed only responded on 69% of occasions (n = 27), their response was almost 4 times more likely to escalate to one of aggression. Our results suggest that certain sites may be preferentially sought after and defended by sea turtles.