Movement patterns for a critically endangered species, the Leatherback Turtle (Dermochelys coriacea), linked to foraging success and population status

Foraging success for pelagic vertebrates may be revealed by horizontal and vertical movement patterns. We show markedly different patterns for leatherback turtles in the North Atlantic versus Eastern Pacific, which feed on gelatinous zooplankton that are only occasionally found in high densities. In the Atlantic, travel speed was characterized by two modes, indicative of high foraging success at low speeds (<15 km d−1) and transit at high speeds (20–45 km d−1). Only a single mode was evident in the Pacific, which occurred at speeds of 21 km d−1 indicative of transit. The mean dive depth was more variable in relation to latitude but closer to the mean annual depth of the thermocline and nutricline for North Atlantic than Eastern Pacific turtles. The most parsimonious explanation for these findings is that Eastern Pacific turtles rarely achieve high foraging success. This is the first support for foraging behaviour differences between populations of this critically endangered species and suggests that longer periods searching for prey may be hindering population recovery in the Pacific while aiding population maintenance in the Atlantic.

Does prey size matter? Novel observations of feeding in the leatherback turtle (Dermochelys coriacea) allow a test of predator–prey size relationships

Optimal foraging models predict that large predators should concentrate on large prey in order to maximize their net gain of energy intake. Here, we show that the largest species of sea turtle, Dermochelys coriacea, does not strictly adhere to this general pattern. Field observations combined with a theoretical model suggest that a 300 kg leatherback turtle would meet its energetic requirements by feeding for 3–4 h a day on 4 g jellyfish, but only if prey were aggregated in high-density patches. Therefore, prey abundance rather than prey size may, in some cases, be the overriding parameter for foraging leatherbacks. This is a classic example where the presence of small prey in the diet of a large marine predator may reflect profitable foraging decisions if the relatively low energy intake per small individual prey is offset by high encounter rates and minimal capture and handling costs. This study provides, to our knowledge, the first quantitative estimates of intake rate for this species.

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.

Do leatherback turtles Dermochelys coriacea forage during the breeding season? A combination of data-logging devices provide new insights

Animals which undertake migrations from foraging grounds to suitable breeding areas must adopt strategies in these new conditions in order to minimise the rate at which body condition deteriorates (which will occur due to oogenesis or provisioning for young). For some animals this involves continuing foraging, whereas for others the optimal strategy is to fast during the breeding season. The leatherback turtle undertakes long-distance migrations from temperate zones to tropical breeding areas, and in some of these areas it has been shown to exhibit diving behaviour indicative of foraging. We used conventional time–depth recorders and a single novel mouth-opening sensor to investigate the foraging behaviour of leatherback turtles in the southern Caribbean. Diving behaviour suggested attempted foraging on vertically migrating prey with significantly more diving to a more consistent depth occurring during the night. No obvious prey manipulation was detected by the mouth sensor, but rhythmic mouth opening did occur during specific phases of dives, suggesting that the turtle was relying on gustatory cues to sense its immediate environment. Patterns of diving in conjunction with these mouth-opening activities imply that leatherbacks are attempting to forage during the breeding season and that gustatory cues are important to leatherbacks.

Global patterns for upper ceilings on migration distance in sea turtles and comparisons with fish, birds and mammals

1. Some animals migrate huge distances in search of resources with locomotory mode (flying/swimming/walking) thought to drive the upper ceilings on migration distance. Yet in cross-taxa comparisons, upper ceilings on migration distance have been ignored for one important group, sea turtles. 2. Using migration distances recorded for 407 adult and 4715 juvenile sea turtles across five species, we show that for adult cheloniid turtles, the upper ceiling on species migration distances between breeding and foraging habitats (1050–2850 km across species) is similar to that predicted for equivalent-sized marine mammals and fish. 3. In contrast, by feeding in the open ocean, adult leatherback turtles (Dermochelys coriacea) and juveniles of all turtle species can travel around 12 000 km from their natal regions, travelling across the widest ocean basins. For juvenile turtles, this puts their maximum migration distances well beyond those expected for equivalent-sized marine mammals and fish, but not those found in some similar sized birds. 4. Post-hatchling turtles perform these long-distance migrations to juvenile foraging sites only once in their lifetime, while adult turtles return to their breeding sites every few (generally ?2) years. Our results highlight the important roles migration periodicity and foraging mode can play in driving the longest migrations, and the implications for Marine Protected Area planning are considered in terms of sea turtle conservation.

Trophic status drives interannual variability in nesting numbers of marine turtles

Large annual fluctuations are seen in breeding numbers in many populations of non–annual breeders. We examined the interannual variation in nesting numbers of populations of green (Chelonia mydas) (n = 16 populations), loggerhead (Caretta caretta) (n =10 populations), leatherback (Dermochelys coriacea) (n = 9 populations) and hawksbill turtles (Eretmochelys imbricata) (n = 10 populations). Interannual variation was greatest in the green turtle. When comparing green and loggerhead turtles nesting in Cyprus we found that green turtles were more likely to change the interval between laying seasons and showed greater variation in the number of clutches laid in a season. We suggest that these differences are driven by the varying trophic statuses of the different species. Green turtles are herbivorous, feeding on sea grasses and macro–algae, and this primary production will be more tightly coupled with prevailing environmental conditions than the carnivorous diet of the loggerhead turtle.

Satellite telemetry suggests high levels of fishing-induced mortality in marine turtles

Long-term records of nesting numbers, or proxies to nesting numbers, show a precipitous decline in the size of many sea turtle populations. Population declines are most frequently attributed to fisheries bycatch, although direct quantification of this level of mortality is rare. We used satellite-tracking records for turtles in the Mediterranean Sea and Pacific, Atlantic and Indian Oceans to identify when turtles had been captured. Evidence for capture came from a combination of an increase in good quality locations from transmitters, transmitters moving inland to coastal towns and villages, and on-board submergence data, showing that transmitters had come out of the water. A high level of mortality was calculated, confirming current concerns regarding the outlook for sea turtles.

Behaviour and buoyancy regulation in the deepest-diving reptile : the leatherback turtle

In the face of the physical and physiological challenges of performing breath-hold deep dives, marine vertebrates have evolved different strategies. Although behavioural strategies in marine mammals and seabirds have been investigated in detail, little is known about the deepest-diving reptile – the leatherback turtle (Dermochelys coriacea). Here, we deployed tri-axial accelerometers on female leatherbacks nesting on St Croix, US Virgin Islands, to explore their diving strategy. Our results show a consistent behavioural pattern within dives among individuals, with an initial period of active swimming at relatively steep descent angles (∼–40 deg), with a stroke frequency of 0.32 Hz, followed by a gliding phase. The depth at which the gliding phase began increased with the maximum depth of the dives. In addition, descent body angles and vertical velocities were higher during deeper dives. Leatherbacks might thus regulate their inspired air-volume according to the intended dive depth, similar to hard-shelled turtles and penguins. During the ascent, turtles actively swam with a stroke frequency of 0.30 Hz but with a low vertical velocity (∼0.40 ms–1) and a low pitch angle (∼+26 deg). Turtles might avoid succumbing to decompression sickness (‘the bends’) by ascending slowly to the surface. In addition, we suggest that the low body temperature of this marine ectotherm compared with that of endotherms might help reduce the risk of bubble formation by increasing the solubility of nitrogen in the blood. This physiological advantage, coupled with several behavioural and physical adaptations, might explain the particular ecological niche the leatherback turtle occupies among marine reptiles.

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.

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.