Behavioural plasticity in a large marine herbivore: contrasting patterns of depth utilisation between two green turtle (Chelonia mydas) populations

We used time-depth recorders to measure depth utilisation in gravid green turtles (Chelonia mydas) during the internesting period at northern Cyprus (Mediterranean), a nesting area where individuals feed, and at Ascension Island (mid-Atlantic), a nesting area where individuals fast. There were contrasting patterns of depth utilisation between the two sites, illustrating that the behaviour of this species is shaped by local conditions. For example, the amount of time spent shallower than 4 m was 90% at Cyprus but only 31% at Ascension Island, and there was a clear difference between the mean depth at Cyprus (2.7 m, n=9 internesting intervals) versus Ascension Island (9.5 m, n=6 internesting intervals) (t 5=5.92, P=0.002). At Cyprus, turtles spent the greatest percentage of their time at very shallow depths, where surveys reveated a high abundance of seagrass on which this population feeds. In contrast, the deeper distribution at Ascension Island may reflect the preferred depth for resting on the seabed.

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–.

Estimating the number of green and loggerhead turtles nesting annually in the Mediterranean

Most species of marine turtle breed every two or more years and it is the norm for females to lay more than one clutch of eggs within a nesting season. Knowing the interval between breeding seasons and the clutch frequency (number of clutches laid by an individual in a breeding season) of females allows us to assess the status of a nesting population. At Alagadi Beach, Northern Cyprus, over a period of 6 years (1995–2000), we attributed 96% of green Chelonia mydas and 80% of loggerhead Caretta caretta turtle clutches to known individual females. This intensive level of monitoring enabled us to estimate the clutch frequency for both species. Using four different methods we estimated clutch frequency to be 2.9–3.1 clutches per female for green turtles and 1.8–2.2 clutches per female for loggerhead turtles. The median interval between nesting seasons for green turtles was 3 years, and for loggerhead turtles it was 2 years. Utilizing these parameters and available data from other beaches that are monitored regularly, we estimate that there are 2,280–2,787 logger-head and 339–360 green turtles nesting annually at these sites in the Mediterranean. This highlights the Critically Endangered status of this population of green turtles. Furthermore, as conventional beach patrols underestimate clutch frequency, these population estimates are likely to be optimistic.

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.

Reproductive seasonality and sexual dimorphism in green turtles

The temporal distribution of nesting and mating in green turtles Chelonia mydas at Ascension Island (7°57¹S, 14°22¹W) in the South Atlantic is described. Mathematical description of the seasonal pattern of nesting showed extreme similarity between seasons, and evidence is presented to support the hypothesis that observed patterns are driven by prevailing environmental temperature. Mating was observed to begin before nesting and follow a pattern consistent with a modelled seasonal influx of suitable females into the annual breeding population. When available data on male size are compared with that of females from the same population (n = 12 populations), a pronounced and consistent sexual dimorphism, with males being smaller than females, is highlighted in all populations. The possible mechanisms behind the evolution of such a pattern are discussed.

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.

Testing the navigational abilities of ocean migrants: displacement experiments on green sea turtles (Chelonia mydas)

Like many animals migrating through the oceans, sea turtles face difficult navigational tasks when they have to reach distant, specific sites. The paradigmatic case of Brazilian green turtles (Chelonia mydas), which nest on the tiny Ascension Island in the middle of the Atlantic Ocean, has often been the subject of hypotheses concerning their navigational mechanisms. To investigate their nature, we displaced 18 females from Ascension and tracked them by satellite after release from eight different points in the ocean, 60–450 km away from the island. Four turtles moved to Brazil soon after the release, 4 moved in various directions before heading to Brazil, and 10 reached the island. All the successful trips, bar 1, were winding but ended with a final straight segment of variable length, as if the turtles were searching for a sensory contact with the island which they obtained at various distances. The approach to Ascension mostly occurred from the direction opposite to the trade wind, suggesting a navigational role of wind-borne information originating from the island.

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.

Dive angles for a green turtle (Chelonia mydas)

The depth and swim speed of a green turtle (Chelonia mydas) were measured during the internesting period in Cyprus. For dives to the seabed (U-dives) we used these data to determine dive angles. Typically the turtle initially descended at a steep angle ([similar]60&deg;) but as the dive continued this angle lessened until the turtle approached the seabed at an average angle of [similar]15&deg;. This systematic change in descent angle is consistent with the prediction that the energetic implications of dive angle are most important at the start of the dive when the turtle is fighting to overcome its positive buoyancy. On leaving the seabed, the turtle often seemed to rise passively.

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&deg;57'S, 14&deg;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&deg;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.