Jellyfish aggregations and leatherback turtle foraging patterns in a temperate coastal environment

Leatherback turtles (Dermochelys coriacea) are obligate predators of gelatinous zooplankton. However, the spatial relationship between predator and prey remains poorly understood beyond sporadic and localized reports. To examine how jellyfish (Phylum Cnidaria: Orders Semaeostomeae and Rhizostomeae) might drive the broad-scale distribution of this wide ranging species, we employed aerial surveys to map jellyfish throughout a temperate coastal shelf area bordering the northeast Atlantic. Previously unknown, consistent aggregations of Rhizostoma octopus extending over tens of square kilometers were identified in distinct coastal “hotspots” during consecutive years (2003–2005). Examination of retrospective sightings data (>50 yr) suggested that 22.5% of leatherback distribution could be explained by these hotspots, with the inference that these coastal features may be sufficiently consistent in space and time to drive long-term foraging associations.

Pan-atlantic analysis of the overlap of a highly migratory species, the leatherback turtle, with pelagic longline fisheries

Large oceanic migrants play important roles in ecosystems, yet many species are of conservation concern as a result of anthropogenic threats, of which incidental capture by fisheries is frequently identified. The last large populations of the leatherback turtle, Dermochelys coriacea, occur in the Atlantic Ocean, but interactions with industrial fisheries could jeopardize recent positive population trends, making bycatch mitigation a priority. Here, we perform the first pan-Atlantic analysis of spatio-temporal distribution of the leatherback turtle and ascertain overlap with longline fishing effort. Data suggest that the Atlantic probably consists of two regional management units: northern and southern (the latter including turtles breeding in South Africa). Although turtles and fisheries show highly diverse distributions, we highlight nine areas of high susceptibility to potential bycatch (four in the northern Atlantic and five in the southern/equatorial Atlantic) that are worthy of further targeted investigation and mitigation. These are reinforced by reports of leatherback bycatch at eight of these sites. International collaborative efforts are needed, especially from nations hosting regions where susceptibility to bycatch is likely to be high within their exclusive economic zone (northern Atlantic: Cape Verde, Gambia, Guinea Bissau, Mauritania, Senegal, Spain, USA and Western Sahara; southern Atlantic: Angola, Brazil, Namibia and UK) and from nations fishing in these high-susceptibility areas, including those located in international waters.

Sand temperatures for nesting sea turtles in the Caribbean: Implications for hatchling sex ratios in the face of climate change

A 200-year time series of incubation temperatures and primary sex ratios for green (Chelonia mydas), hawksbill (Eretmochelys imbricata) and leatherback (Dermochelys coriacea) sea turtles nesting in St. Eustatius (North East Caribbean) was created by combining sand temperature measurements with historical and current environmental data and climate projections. Rainfall and spring tides were important because they cooled the sand and lowered incubation temperatures. Mean annual sand temperatures are currently 31.0. °C (SD. =. 1.6) at the nesting beach but show seasonality, with lower temperatures (29.1-29.6. °C) during January-March and warmer temperatures (31.9-33.3. °C) in June-August. Results suggest that all three species have had female-biased hatchling production for the past decades with less than 15.5%, 36.0%, and 23.7% males produced every year for greens, hawksbills and leatherbacks respectively since the late nineteenth century. Global warming will exacerbate this female-skew. For example, projections indicate that only 2.4% of green turtle hatchlings will be males by 2030, 1.0% by 2060, and 0.4% by 2090. On the other hand, future changes to nesting phenology have the potential to mitigate the extent of feminisation. In the absence of such phenological changes, management strategies to artificially lower incubation temperatures by shading nests or relocating nest clutches to deeper depths may be the only way to prevent the localised extinction of these turtle populations.

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.

Movements of migrating green turtles in relation to AVHRR derived sea surface temperature

Previous studies have shown that for some populations of marine turtle, individuals move along narrow migration corridors in the open ocean. It has been suggested that these migration corridors may correspond with nearsurface oceanographic features that can be detected by remote sensing. This idea is examined by superimposing the tracks of green turtles (Chelonia mydas) migrating from Ascension Island to Brazil, on sea surface temperature (SST) data derived from Advanced Very High Resolution Radiometer (AVHRR) images. The turtles did not follow specific isotherms during migration nor make turns en-route where specific thermal cues were encountered. These results suggest that for this population, SST plays a minimal role in influencing the exact route that individuals follow.

Life in the really slow lane : loggerhead sea turtles mature late relative to other reptiles

1. Age at maturity is hard to estimate for species that cannot be directly marked or observed throughout their lives and yet is a key demographic parameter that is needed to assess the conservation status of endangered species. 2. For loggerhead turtles (Caretta caretta) in the North Atlantic and North Pacific, juvenile growth rates (c. 10 cm year−1) were calculated by examining size increases during transoceanic journeys; durations of which were estimated from satellite-tracked Lagrangian surface drifter buoy trajectories. 3. Lagrangian-derived growth estimates were used in a weighted loglinear model of size-specific growth rates for loggerhead turtles and combined with newly available information on size at maturity to estimate an age at maturity of 45 years (older than past estimates). 4. By examining the age at maturity for 79 reptile species, we show that loggerhead turtles, along with other large-bodied Testudine (turtle and tortoise) species, take longer to reach maturity than other reptile species of comparable sizes. This finding heightens concern over the future sustainability of turtle populations. By maturing at an old age, sea turtles will be less resilient to anthropogenic mortality than previously suspected.

Reproductive investment and optimum clutch size of Loggerhead Sea Turtles (Caretta caretta)

(1) Data for loggerhead turtles (Caretta caretta L.) nesting on the Greek island of Cephalonia were used to develop a model which predicted the optimum clutch size. (2) There was a positive linear relationship between the number of clutches into which eggs could be divided and the total time spent by a nesting turtle on the beach, and hence a negative relationship between the time invested on the beach per egg and clutch size. (3) A previous study indicated that energy expenditure for nesting turtles on land is very high, so there may be a selective pressure to maximize clutch size in order to minimize the energy expended per egg laid. As there appeared to be no counterselective pressures favouring small clutches, clutch size should be constrained by a female's egg-carryingc apacity,w hich in turn could be expected to be related to her body size. Hence, a positive relationship between clutch size and body size was predicted, and was found in the population under study.

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.

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.

Genetic composition of the Ascension Island green turtle rookery based on mitochondrial DNA : implications for sampling and diversity

In species of conservation concern it is often difficult to be certain that population diversity and structure have been adequately characterised by genetic sampling. Since practical and financial constraints tend to be associated with increasing sample sizes in many conservation genetic studies, it is important to consider the potential for sampling error and bias due to inadequate samples or spatio-temporal structure within populations. We analysed sequence data from the mitochondrial DNA control region in a large sample (n = 245) of green sea turtles Chelonia mydas collected at the globally important rookery of Ascension Island, South Atlantic. We examined genetic diversity and structure among 10 sampling sites, 4 beach clusters and 4 nesting seasons, and evaluated the genetic composition of Ascension against other Atlantic nesting populations, including the well-studied rookery at Tortuguero (Costa Rica). Finally, we used rarefaction and GENESAMP analyses to assess the ability of different sample sizes to provide acceptable genetic representations of a population, using Ascension and Tortuguero as models. On Ascension, we found 13 haplotypes, of which only 3 had been previously observed in the rookery, and 5 previously undescribed. We detected no differentiation among beach clusters or sampling seasons, and only weak differentiation among the 3 primary nesting sites. The increased sample size for Ascension provided higher resolution and statistical power in describing genetic structure among all other known Atlantic rookeries. Our extrapolations showed that a maximum of 18 and 6 haplotypes are expected to occur in Ascension and Tortuguero, respectively, and that current sample sizes are sufficient to describe most of the variation. We recommend using rarefaction and GENESAMP analyses on a rookery-by-rookery basis to evaluate whether a sample set adequately describes mitochondrial DNA diversity, thus strengthening subsequent phylogeographic and mixed stock analyses, and management recommendations for conservation.

Vertical movements of North Sea Cod

Various air-breathing marine vertebrates such as seals, turtles and seabirds show distinct patterns of diving behaviour. For fish, the distinction between different vertical behaviours is often less clear-cut, as there are no surface intervals to differentiate between dives. Using data from acoustic tags (n = 23) and archival depth recorders attached to cod Gadus morhua (n = 92) in the southern North Sea, we developed a quantitative method of classifying vertical movements in order to facilitate an objective comparison of the behaviour of different individuals. This method expands the utilisation of data from data storage tags, with the potential for a better understanding of fish behaviour and enhanced individual based behaviour for improved ecosystem modelling. We found that cod were closely associated with the seabed for 90% of the time, although they showed distinct seasonal and spatial patterns in behaviour. For example, cod tagged in the southern North Sea exhibited high rates of vertical movement in spring and autumn that were probably associated with migration, while the vertical movements of resident cod in other areas were much less extensive and were probably related to foraging or spawning behaviours. The full reasons underlying spatial and temporal behavioural plasticity by cod in the North Sea warrant further investigation.

Satellite tracking reveals unusual diving characteristics for a marine reptile, the olive ridley turtle : Lepidochelys olivacea

The movements, diving behaviour and thermal environment occupied by 4 adult female olive ridley turtles Lepidochelys olivacea in northern Australia were determined through satellite telemetry. Patterns of behaviour recorded were rather unusual compared to other sea turtles in that dives were mainly deep, largely benthic and exceptionally long (>2 h) in some cases, characteristics typical of over-wintering turtles in colder environments. One individual occupied shallow coastal foraging zones, while the others foraged far from land (probably on the seabed) in relatively deep water (>100 m). Individuals performed long dives (frequently >100 min), but from the short post-dive intervals we suggest that these dives were mainly aerobic. Maximum dive depth recorded was 200 ± 20 m (mean maximum depths ranged from 20.1 to 46.7 m across individuals; n = 17328 dives in total; depths ≥3 m were considered ‘dives’) and the maximum duration was 200 ± 20 min (mean durations ranged from 24.5 to 48.0 min across individuals). Temperature profiles indicate that turtles experienced temperatures ranging from 23 to 29°C at the surface, with the lowest temperature recorded (18.7°C) at a depth of 98 m. Only 6.9% of the dives were in water <20°C. From time-allocation at depth (TAD) scores, we demonstrated that many dives reaching the known or inferred sea bottom were U-shaped, but there was no apparent diel signal in dive depth. This suggests that many benthic dives were not associated exclusively with resting behaviour and likely had a foraging component as well. The ability to perform long benthic dives allows this species to exploit deeper benthic environments in addition to the shallow coastal areas more generally occupied by adult hard-shelled sea turtles (e.g. green and hawksbill turtles). Deep benthic dives also occur in certain marine mammals (e.g. narwhals) and sea birds (e.g. rockhopper penguins) and therefore seem to be a general foraging strategy exploited by animals that can perform long dives.

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.