Migration cues and timing in leatherback sea turtles

Atlantic leatherback sea turtles migrate annually from foraginggrounds off eastern Canada and the northeastern United Statesto southern foraging and breeding areas. Using Cox's proportionalhazards model, we investigated the individual timing of thesouthward migrations of 27 turtles equipped with satellite-linkedtransmitters off Nova Scotia compared with turtle characteristicsand satellite-measured ocean variables. Latitude, longitude,1-week lagged average sea surface temperature, and 1-week laggedaverage chlorophyll-a concentration appear to influence theprobability of departure. Higher temperature and, in the northernrange of the study, higher chlorophyll concentration increaseddeparture rates, perhaps due to the acceleration of the lifecycle of the leatherback's gelatinous prey and/or increasedfeeding efficiency in these areas. This study highlights theopportunity to use satellite telemetry and environmental datato examine the cues for and timing of animal migrations andexpands the study of migration timing to include a new speciesand environment.     

Orientation behaviour of leatherback sea turtles within the North Atlantic subtropical gyre

Leatherback sea turtles (Dermochelys coriacea) travel thousands of kilometres between temperate feeding and tropical breeding/over-wintering grounds, with adult turtles able to pinpoint specific nesting beaches after multi-year absences. Their extensive migrations often occur in oceanic habitat where limited known sensory information is available to aid in orientation. Here, we examined the migratory orientation of adult male, adult female and subadult leatherbacks during their open-ocean movements within the North Atlantic subtropical gyre by analysing satellite-derived tracks from fifteen individuals over a 2-year period. To determine the turtles'' true headings, we corrected the reconstructed tracks for current drift and found negligible differences between current-corrected and observed tracks within the gyre. Individual leatherback headings were remarkably consistent throughout the subtropical gyre, with turtles significantly oriented to the south-southeast. Adult leatherbacks of both sexes maintained similar mean headings and showed greater orientation precision overall. The consistent headings maintained by adult and subadult leatherbacks within the gyre suggest use of a common compass sense.     

Identification of high-use habitat and threats to leatherback sea turtles in northern waters: new directions for conservation

Incidental capture in fisheries threatens many marine vertebrates, however, conservation cannot be effective without identifying major sources of mortality. For the critically endangered leatherback turtle (Dermochelys coriacea), a reliance on fisheries observer data and an absence of behavioural data sets corresponding to a large and diverse sample of animals have focused conservation efforts on a very limited part of the species marine habitat. Using the largest satellite telemetry data set for Atlantic leatherbacks, morphometrics from foraging animals and entanglement records, we show annual return migrations to key feeding areas by males, females and juveniles, and demonstrate the importance of northern latitudes to leatherbacks. We show that leatherbacks are vulnerable to entanglement in northern coastal and shelf waters, where turtle–fishery interactions represent a greater threat to this species than previously recognized. Unless conservation efforts expand to coastal and shelf areas, present efforts alone will not be sufficient to save the species.     

Long-term monitoring of leatherback turtle diving behaviour during oceanic movements

The diving behaviour of four leatherback turtles (Dermochelys coriacea) was recorded for periods of 0.5-8.1 months during their postnesting movements in the Indian and Atlantic Oceans, when they covered 1569-18,994 km. Dive data were obtained using satellite-linked transmitters which also provided information on the dive depths and profiles of the turtles. Turtles mainly dove to depths < 200 m, with maximum dive durations under 30-40 min and exhibited diel variations in their diving activity for most part of the routes, with dives being usually longer at night. Diurnal dives were in general quite short, but cases of very deep (> 900 m) and prolonged (> 70 min) dives were however recorded only during daytime. The three turtles that were tracked for the longest time showed a marked change in behaviour during the tracking, decreasing their dive durations and ceasing to dive deeply. Moreover, diel variations disappeared, with nocturnal dives becoming short and numerous. This change in turtle diving activity appeared to be related to water temperature, suggesting an influence of seasonal prey availability on their diving behaviour. The turtle diving activity was independent on the shape of their routes, with no changes between linear movements in the core of main currents or looping segments in presence of oceanic eddies.     

Current transport of leatherback sea turtles (Dermochelys coriacea) in the ocean

While the long-distance movements of pelagic vertebrates are becoming known thanks to satellite telemetry, the factors determining their courses have hardly been investigated. We have analysed the effects of oceanographic factors on the post-nesting movements of three satellite-tracked leatherback turtles (Dermochelys coriacea) moving in the southwest Indian Ocean. By superimposing the turtle tracks on contemporaneous images of sea-surface temperatures and sea height anomalies, we show that currentrelated features dominate the shape of the reconstructed routes. After an initial offshore movement, turtles moved along straight routes when in the core of the current, or executed loops within eddies. Large parts of the routes were strikingly similar to those of surface drifters tracked in the same region. These findings document that long-lasting oceanic movements of marine turtles may be shaped by oceanic currents.     

Behaviour of leatherback sea turtles, Dermochelys coriacea, during the migratory cycle

Leatherback sea turtles, Dermochelys coriacea, undertake broad oceanic movements. While satellite telemetry has been used to investigate the post-nesting behaviour of female turtles tagged on tropical nesting beaches, long-term behavioural patterns of turtles of different sexes and sizes have not been described. Here we investigate behaviour for 25 subadult and adult male and female turtles satellite-tagged in temperate waters off Nova Scotia, Canada. Although sex and reproductive condition contributed to variation in migratory patterns, the migratory cycle of all turtles included movement between temperate and tropical waters. Marked changes in rates of travel, and diving and surfacing behaviour, accompanied southward movement away from northern foraging areas. As turtles approached higher latitudes the following spring and summer, they assumed behaviours consistent with regular foraging activity and eventually settled in coastal areas off Canada and the northeastern USA. Behavioural patterns corresponding to various phases of the migratory cycle were consistent across multiple animals and were repeated within individuals that completed return movements to northern waters. We consider the potential biological significance of these patterns, including how turtle behaviour relates to predator avoidance, thermoregulation and prey distribution.     

Predicting bycatch hotspots for endangered leatherback turtles on longlines in the Pacific Ocean

Fisheries bycatch is a critical source of mortality for rapidly declining populations of leatherback turtles, Dermochelys coriacea. We integrated use-intensity distributions for 135 satellite-tracked adult turtles with longline fishing effort to estimate predicted bycatch risk over space and time in the Pacific Ocean. Areas of predicted bycatch risk did not overlap for eastern and western Pacific nesting populations, warranting their consideration as distinct management units with respect to fisheries bycatch. For western Pacific nesting populations, we identified several areas of high risk in the north and central Pacific, but greatest risk was adjacent to primary nesting beaches in tropical seas of Indo-Pacific islands, largely confined to several exclusive economic zones under the jurisdiction of national authorities. For eastern Pacific nesting populations, we identified moderate risk associated with migrations to nesting beaches, but the greatest risk was in the South Pacific Gyre, a broad pelagic zone outside national waters where management is currently lacking and may prove difficult to implement. Efforts should focus on these predicted hotspots to develop more targeted management approaches to alleviate leatherback bycatch.     

Evidence for transoceanic migrations by loggerhead sea turtles in the southern Pacific Ocean

Post-hatchling loggerhead turtles (Caretta caretta) in the northern Pacific and northern Atlantic Oceans undertake transoceanic developmental migrations. Similar migratory behaviour is hypothesized in the South Pacific Ocean as post-hatchling loggerhead turtles are observed in Peruvian fisheries, yet no loggerhead rookeries occur along the coast of South America. This hypothesis was supported by analyses of the size-class distribution of 123 post-hatchling turtles in the South Pacific and genetic analysis of mtDNA haplotypes of 103 nesting females in the southwest Pacific, 19 post-hatchlings stranded on the southeastern Australian beaches and 22 post-hatchlings caught by Peruvian longline fisheries. Only two haplotypes (CCP1 93% and CCP5 7%) were observed across all samples, and there were no significant differences in haplotype frequencies between the southwest Pacific rookeries and the post-hatchlings. By contrast, the predominant CCP1 haplotype is rarely observed in North Pacific rookeries and haplotype frequencies were strongly differentiated between the two regions (F_st=0.82; p=<0.00001). These results suggest that post-hatchling loggerhead turtles emerging from the southwest Pacific rookeries are undertaking transoceanic migrations to the southeastern Pacific Ocean, thus emphasizing the need for a broader focus on juvenile mortality throughout the South Pacific to develop effective conservation strategies.     

Tracking leatherback turtles from the world's largest rookery: assessing threats across the South Atlantic

Despite extensive work carried out on leatherback turtles (Dermochelys coriacea) in the North Atlantic and Indo-Pacific, very little is known of the at-sea distribution of this species in the South Atlantic, where the world's largest population nests in Gabon (central Africa). This paucity of data is of marked concern given the pace of industrialization in fisheries with demonstrable marine turtle bycatch in African/Latin American waters. We tracked the movements of 25 adult female leatherback turtles obtaining a range of fundamental and applied insights, including indications for methodological advancement. Individuals could be assigned to one of three dispersal strategies, moving to (i) habitats of the equatorial Atlantic, (ii) temperate habitats off South America or (iii) temperate habitats off southern Africa. While occupying regions with high surface chlorophyll concentrations, these strategies exposed turtles to some of the world's highest levels of longline fishing effort, in addition to areas with coastal gillnet fisheries. Satellite tracking highlighted that at least 11 nations should be involved in the conservation of this species in addition to those with distant fishing fleets. The majority of tracking days were, however, spent in the high seas, where effective implementation of conservation efforts is complex to achieve.     

Quantifying the effects of fisheries on threatened species: the impact of pelagic longlines on loggerhead and leatherback sea turtles

The depletion of fish stocks from global fisheries has been a long‐standing concern. More recently, incidental catch of non‐target (termed bycatch) vertebrates also has been proposed as a serious conservation issue. Here we present a bycatch assessment for loggerhead and leatherback sea turtles that are incidentally caught by global pelagic longlines. We integrate catch data from over 40 nations and bycatch data from 13 international observer programmes. Despite infrequent rates of encounter, our analyses show that more than 200 000 loggerheads and 50 000 leatherbacks were likely taken as pelagic longline bycatch in 2000. Our analyses suggest that thousands of these turtles die each year from longline gear in the Pacific Ocean alone. Given 80–95% declines for Pacific loggerhead and leatherback populations over the last 20 years, this bycatch level is not sustainable. Adopting a large‐scale, synthetic approach is critical to accurately characterize the influence of global fisheries bycatch on globally distributed and imperilled pelagic vertebrates.     

GPS tracking of the foraging movements of Manx Shearwaters Puffinus puffinus breeding on Skomer Island, Wales

We report the first successful use of miniature Global Positioning System loggers to track the ocean‐going behaviour of a c. 400 g seabird, the Manx Shearwater Puffinus puffinus. Breeding birds were tracked over three field seasons during the incubation and chick‐rearing periods on their foraging excursions from the large colony on Skomer Island, Pembrokeshire, UK. Foraging effort was concentrated in the Irish Sea. Likely foraging areas were identified to the north, and more diffusely to the west of the colony. No foraging excursions were recorded significantly to the south of the colony, conflicting with the conclusions of earlier studies based on ringing recoveries and observations. We discuss several explanations including the hypothesis that foraging may have shifted substantially northwards in recent decades. We found no obvious relationship between birds’ positions and water depth, although there was a suggestion that observations at night were in shallower water than those during the day. We also found that, despite the fact that Shearwaters can be observed rafting off‐shore from their colonies in the hours prior to making landfall at night, breeding birds are usually located much further from the colony in the last 8 h before arrival, a finding that has significance for the likely effectiveness of marine protection areas if they are only local to the colony. Short sequences of precise second‐by‐second fixes showed that movement speeds were bimodal, corresponding to sitting on the water (most common at night and around midday) and flying (most common in the morning and evening), with flight behaviour separable into erratic (indicative of searching for food) and directional (indicative of travelling). We also provide a first direct measurement of mean flight speed during directional flight (c. 40 km/h), slower than a Shearwater's predicted maximum range velocity, suggesting that birds are exploiting wave or dynamic soaring during long‐distance travel.     

Foraging behavior of adult female Steller sea lions during the breeding season in Southeast Alaska

During the 1990s, the Steller sea lion ( Eumetopias jubatus Schreber) Western Alaska stock (WS) suffered steep population decline while the Eastern Alaska stock (ES) steadily increased. One bottom-up forcing hypothesis explaining this decline predicted lactating adult female foraging behavior would be different between stocks. To investigate this effect, we monitored 11 ES females at two breeding rookeries using satellite dive recorders (SDR) during the early breeding seasons of 1992–1993, examined their behavior with respect to prey, physiological limitations, and habitat, and made limited comparisons to observations of WS female behavior reported in the literature. ES females were not operating at the extremes of ability, with most diving within the limits of aerobic metabolism, less than one-quarter of possible foraging time during trips spent submerged and most foraging trips requiring less than one-half the lipid store fasting ability of dependent pups. Thus, females may have some capacity to alter behavior to accommodate future changes in foraging conditions, but the extent of this plasticity is unknown. Because recent work suggests WS recovery is impeded by low natality, future studies should test differences between reproductive and non-reproductive mature females in order to properly assess the contribution of foraging ecology to SSL population dynamics.     

Topsy-turvy: turning the counter-current heat exchange of leatherback turtles upside down

Counter-current heat exchangers associated with appendages of endotherms feature bundles of closely applied arteriovenous vessels. The accepted paradigm is that heat from warm arterial blood travelling into the appendage crosses into cool venous blood returning to the body. High core temperature is maintained, but the appendage functions at low temperature. Leatherback turtles have elevated core temperatures in cold seawater and arteriovenous plexuses at the roots of all four limbs. We demonstrate that plexuses of the hindlimbs are situated wholly within the hip musculature, and that, at the distal ends of the plexuses, most blood vessels supply or drain the hip muscles, with little distal vascular supply to, or drainage from the limb blades. Venous blood entering a plexus will therefore be drained from active locomotory muscles that are overlaid by thick blubber when the adults are foraging in cold temperate waters. Plexuses maintain high limb muscle temperature and avoid excessive loss of heat to the core, the reverse of the accepted paradigm. Plexuses protect the core from overheating generated by muscular thermogenesis during nesting.     

High juvenile mortality during migration in a declining population of a long‐distance migratory raptor

Many populations of long‐distance migrants are declining and there is increasing evidence that declines may be caused by factors operating outside the breeding season. Among the four vulture species breeding in the western Palaearctic, the species showing the steepest population decline, the Egyptian Vulture Neophron percnopterus, is a long‐distance migrant wintering in Africa. However, the flyways and wintering areas of the species are only known for some populations, and without knowledge of where mortality occurs, effective conservation management is not possible. We tracked 19 juvenile Egyptian Vultures from the declining breeding population on the Balkan Peninsula between 2010 and 2014 to estimate survival and identify important migratory routes and wintering areas for this species. Mortality during the first autumn migration was high (monthly survival probability 0.75) but mortality during migration was exclusively associated with suboptimal navigation. All birds from western breeding areas and three birds from central and eastern breeding areas attempted to fly south over the Mediterranean Sea, but only one in 10 birds survived this route, probably due to stronger tailwind. All eight birds using the migratory route via Turkey and the Middle East successfully completed their first autumn migration. Of 14 individual and environmental variables examined to explain why juvenile birds did or did not successfully complete their first migration, the natal origin of the bird was the most influential. We speculate that in a declining population with fewer experienced adults, an increasing proportion of juvenile birds are forced to migrate without conspecific guidance, leading to high mortality as a consequence of following sub‐optimal migratory routes. Juvenile Egyptian Vultures wintered across a vast range of the Sahel and eastern Africa, and had large movement ranges with core use areas at intermediate elevations in savannah, cropland or desert. Two birds were shot in Africa, where several significant threats exist for vultures at continental scales. Given the broad distribution of the birds and threats, effective conservation in Africa will be challenging and will require long‐term investment. We recommend that in the short term, more efficient conservation could target narrow migration corridors in southern Turkey and the Middle East, and known congregation sites in African wintering areas.     

Life-cycle strategies and seasonal migrations of oceanic copepods in the Irminger Sea

Abundance and seasonal vertical distribution of dominant zooplankters in the Irminger Sea was studied from data collected during four cruises between November 1996 and June 1997. In addition, egg production of Calanus finmarchicus was measured during winter, spring and summer 1996–2001. Five taxa constituted >95% of the copepod biomass, C. finmarchicus, Pareuchaeta norvegica, C. hyperboreus, Oithona spp. and Oncaea spp. A seasonal migration pattern was evident for C. finmarchicus, P. norvegica and Oithona spp.: from December to February, they inhabited the deeper layers, whereas, from April to June, they were most abundant in the upper layers. Oncaea spp. also stayed deep during winter and only a very limited part of the population rose to the surface during summer. C. hyperboreus remained deep from December to April, but had virtually disappeared in June. Reproduction of C. finmarchicus took place in May in the surface layers and was linked to the phytoplankton spring bloom. In contrast, reproduction of P. norvegica occurred at depth in February and was uncoupled with the spring bloom. C. hyperboreus did not reproduce in the Irminger Sea. Data on Oithona spp. and Oncaea spp. indicated that the former reproduced between April and June in the upper layers, whereas the latter reproduced year-round at depth. Thus, data on vertical distribution and seasonal stage composition suggested that the dominant copepods are separated, at least partly, at spatial and temporal scales with regard to overwintering and development.     

Robust hierarchical state-space models reveal diel variation in travel rates of migrating leatherback turtles

1. Biological and statistical complexity are features common to most ecological data that hinder our ability to extract meaningful patterns using conventional tools. Recent work on implementing modern statistical methods for analysis of such ecological data has focused primarily on population dynamics but other types of data, such as animal movement pathways obtained from satellite telemetry, can also benefit from the application of modern statistical tools. 2. We develop a robust hierarchical state-space approach for analysis of multiple satellite telemetry pathways obtained via the Argos system. State-space models are time-series methods that allow unobserved states and biological parameters to be estimated from data observed with error. We show that the approach can reveal important patterns in complex, noisy data where conventional methods cannot. 3. Using the largest Atlantic satellite telemetry data set for critically endangered leatherback turtles, we show that the diel pattern in travel rates of these turtles changes over different phases of their migratory cycle. While foraging in northern waters the turtles show similar travel rates during day and night, but on their southward migration to tropical waters travel rates are markedly faster during the day. These patterns are generally consistent with diving data, and may be related to changes in foraging behaviour. Interestingly, individuals that migrate southward to breed generally show higher daytime travel rates than individuals that migrate southward in a non-breeding year. 4. Our approach is extremely flexible and can be applied to many ecological analyses that use complex, sequential data.     

Habitat and behaviour of yellowfin tuna Thunnus albacares in the Gulf of Mexico determined using pop-up satellite archival tags

This study presents the first data on movement, habitat use and behaviour for yellowfin tuna Thunnus albacares in the Atlantic Basin. Six individuals were tracked in the Gulf of Mexico using pop-up satellite archival tags. Records up to 80 days in length were obtained, providing information on depth and temperature preferences as well as horizontal movements. Thunnus albacares in the Gulf of Mexico showed a strong preference for the mixed layer and thermocline, consistent with findings for this species in other ocean basins. Fish showed a diel pattern in depth distribution, remaining in surface and mixed layer waters at night and diving to deeper waters during the day. The vertical extent of T. albacares habitat appeared to be temperature limited, with fish generally avoiding waters that were >6° C cooler than surface waters. The vertical and thermal habitat usage of T. albacares differs from that of bigeye Thunnus obesus and bluefin Thunnus thynnus , Thunnus orientalis and Thunnus maccoyii tunas. These results are consistent with the results of earlier studies conducted on T. albacares in other oceans.     

Temperature and the life-history strategies of sea turtles

1. 1. Sea turtles have a high fecundity, high mortality, great longevity life history strategy.

2. 2. With the exception of the leatherback, turtle distribution is constrained by the 20°C surface isotherm.

3. 3. All sea turtles exhibit temperature-dependent sex determination (TSD) with pivotal temperatures close to 29°C.

4. 4. It is suggested that hatchling sex ratio will vary chaotically because of TSD.

5. 5. Because of TSD and natal homing, sea turtles are likely to be adversely affected by global warming.

6. 6. TSD and global warming have implications for conservation/management of sea turtles.