Spatial variation of plastic debris on important turtle nesting beaches of the remote Chagos Archipelago, Indian Ocean.

We report Anthropogenic Marine Debris (AMD) in Chagos Archipelago in the Indian Ocean, globally amongst the most isolated island groups. AMD on 14 island beaches in five atolls were surveyed in 2019 using two techniques: Marine Debris Tracker (MDT) along littoral vegetation and photoquadrats in open beach. Over 60 % of AMD in both beach zones was composed of plastics, especially bottles and fragments (mean = 44.9 %, 27.2 %, range = 16.5-73.2 %, 4.8-55.9 % respectively in vegetation; mean = 28.7 %, 31.5 %, range = 17.7-40.7 %, 11.6-60.0 % respectively in open beach). The density of plastic debris in littoral vegetation (MDT data: 1995 bottles, 3328 fragments per 100 m2) was 10-fold greater than in open beach (photoquadrat data: 184 bottles, 106 fragments per 100 m2). Significant latitudinal variation in vegetation AMD occurred (8-fold greater in southern atolls, p = 0.006). AMD varied within island zones: most debris observed on oceanside beaches (oceanside vs lagoon, W = 365, p < 0.001; ocean vs island tip, W = 107, p = 0.034). Standardisation of surveys using the open-source MDT App is recommended. Debris accumulation hotspots overlapped with sea turtle nesting habitat, guiding future beach clean-up prioritisation.

The importance of sample size in marine megafauna tagging studies.

Telemetry is a key, widely used tool to understand marine megafauna distribution, habitat use, behavior, and physiology; however, a critical question remains: "How many animals should be tracked to acquire meaningful data sets?" This question has wide-ranging implications including considerations of statistical power, animal ethics, logistics, and cost. While power analyses can inform sample sizes needed for statistical significance, they require some initial data inputs that are often unavailable. To inform the planning of telemetry and biologging studies of marine megafauna where few or no data are available or where resources are limited, we reviewed the types of information that have been obtained in previously published studies using different sample sizes. We considered sample sizes from one to >100 individuals and synthesized empirical findings, detailing the information that can be gathered with increasing sample sizes. We complement this review with simulations, using real data, to show the impact of sample size when trying to address various research questions in movement ecology of marine megafauna. We also highlight the value of collaborative, synthetic studies to enhance sample sizes and broaden the range, scale, and scope of questions that can be answered.

The discovery of deep-water seagrass meadows in a pristine Indian Ocean wilderness revealed by tracking green turtles.

Our understanding of global seagrass ecosystems comes largely from regions characterized by human impacts with limited data from habitats defined as notionally pristine. Seagrass assessments also largely focus on shallow-water coastal habitats with comparatively few studies on offshore deep-water seagrasses. We satellite tracked green turtles (Chelonia mydas), which are known to forage on seagrasses, to a remote, pristine deep-water environment in the Western Indian Ocean, the Great Chagos Bank, which lies in the heart of one of the world's largest marine protected areas (MPAs). Subsequently we used in-situ SCUBA and baited video surveys to survey the day-time sites occupied by turtles and discovered extensive monospecific seagrass meadows of Thalassodendron ciliatum. At three sites that extended over 128 km, mean seagrass cover was 74% (mean range 67-88% across the 3 sites at depths to 29 m. The mean species richness of fish in seagrass meadows was 11 species per site (mean range 8-14 across the 3 sites). High fish abundance (e.g. Siganus sutor: mean MaxN.site-1 = 38.0, SD = 53.7, n = 5) and large predatory shark (Carcharhinus amblyrhynchos) (mean MaxN.site-1 = 1.5, SD = 0.4, n = 5) were recorded at all sites. Such observations of seagrass meadows with large top predators, are limited in the literature. Given that the Great Chagos Bank extends over approximately 12,500 km2 and many other large deep submerged banks exist across the world's oceans, our results suggest that deep-water seagrass may be far more abundant than previously suspected.

Convergence of marine megafauna movement patterns in coastal and open oceans.

The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals' movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyze a global dataset of ∼2.8 million locations from >2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim, walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they move through, resulting in complex movement patterns when moving close to coasts compared with more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal microhabitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise, and declining oxygen content.

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.

When surfacers do not dive: multiple significance of extended surface times in marine turtles.

Marine turtles spend more than 90% of their life underwater and have been termed surfacers as opposed to divers. Nonetheless turtles have been reported occasionally to float motionless at the surface but the reasons for this behaviour are not clear. We investigated the location, timing and duration of extended surface times (ESTs) in 10 free-ranging loggerhead turtles (Caretta caretta) and the possible relationship to water temperature and diving activity recorded via satellite relay data loggers for 101-450 days. For one turtle that dived only in offshore areas, ESTs contributed 12% of the time whereas for the other turtles ESTs contributed 0.4-1.8% of the time. ESTs lasted on average 90 min but were mostly infrequent and irregular, excluding the involvement of a fundamental regulatory function. However, 82% of the ESTs occurred during daylight, mostly around noon, suggesting a dependence on solar radiation. For three turtles, there was an appreciable (7 degrees C to 10.5 degrees C) temperature decrease with depth for dives during periods when ESTs occurred frequently, suggesting a re-warming function of EST to compensate for decreased body temperatures, possibly to enhance digestive efficiency. A positive correlation between body mass and EST duration supported this explanation. By contrast, night-active turtles that exceeded their calculated aerobic dive limits in 7.6-16% of the dives engaged in nocturnal ESTs, probably for lactate clearance. This is the first evidence that loggerhead turtles may refrain from diving for at least two reasons, either to absorb solar radiation or to recover from anaerobic activity.

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.

Asynchronous emergence by loggerhead turtle (Caretta caretta) hatchlings.

For many decades it has been accepted that marine turtle hatchlings from the same nest generally emerge from the sand together. However, for loggerhead turtles (Caretta caretta) nesting on the Greek Island of Kefalonia, a more asynchronous pattern of emergence has been documented. By placing temperature loggers at the top and bottom of nests laid on Kefalonia during 1998, we examined whether this asynchronous emergence was related to the thermal conditions within nests. Pronounced thermal variation existed not only between, but also within, individual nests. These within-nest temperature differences were related to the patterns of hatchling emergence, with hatchlings from nests displaying large thermal ranges emerging over a longer time-scale than those characterised by more uniform temperatures. In many egg-laying animals, parental care of the offspring may continue while the eggs are incubating and also after they have hatched. Consequently, the importance of the nest site for determining incubation conditions may be reduced since the parents themselves may alter the local environment. By contrast, in marine turtles, parental care ceases once the eggs have been laid and the nest site covered. The positioning of the nest site, in both space and time, may therefore have profound effects for marine turtles by affecting, for example, the survival of the eggs and hatchlings as well as their sex (Janzen and Paukstis 1991). During incubation, sea turtle embryos grow from a few cells at oviposition to a self-sufficient organism at hatching some 50-80 days later (Ackerman 1997). After hatching, the young turtles dig up through the sand and emerge typically en masse at the surface 1-7 nights later, with a number of stragglers following over the next few nights (Christens 1990). This contrasts with the frequently observed pattern of hatching asynchrony in birds. It has been suggested that the cause of mass emergence in turtles is that eggs within a clutch are fertilised within a short period of time and then, when thermal conditions within the nest are uniform, develop at very similar rates and hence hatch and emerge together (Porter 1972). As a corollary of this idea, it would be predicted that when there are pronounced within-nest thermal gradients, development rates of siblings will be different and hence asynchronous hatching and emergence might occur. While it may be energetically beneficial for hatchlings to emerge in a group (Carr and Hirth 1961), if the extent of hatching asynchrony is marked then there may be severe costs for individuals if they wait for all their siblings to hatch before attempting to dig out of the sand (Hays and Speakman 1992). Under such conditions, the protracted emergence of small groups of hatchlings over several nights may be favoured. Examination of the literature suggests that emergence asynchrony may be more widespread than generally considered. For example, Witherington et al. (1990) described loggerhead turtle hatchlings (Caretta caretta) emerging over 4 days in Florida; for green turtles (Chelonia mydas), Hendrickson (1958) documented that nests in Malaysia and Sarawak produced hatchlings for up to 8 days; whilst Diamond (1976) found that hawksbill (Eretmochelys imbricata) nests on Cousin Island, Seychelles, were active for up to 4 days. Similarly, on the Greek Island of Kefalonia, we have shown that emergence from individual loggerhead turtle nests may occur on up to 11 nights (Hays and Speakman 1992). It is logical to suppose that asynchronous emergence relates to thermal gradients within nests, since the incubation duration of sea turtle eggs is related to temperature, with eggs hatching quicker when the temperature is higher. Here we test this hypothesis by measuring thermal variations within loggerhead turtle nests and comparing these variations to the patterns of hatchling emergence.

Metabolic heating and the prediction of sex ratios for green turtles (Chelonia mydas).

We compared incubation temperatures in nests (n=32) of the green turtle (Chelonia mydas) on Ascension Island in relation to sand temperatures of control sites at nest depth. Intrabeach thermal variation was low, whereas interbeach thermal variation was high in both control and nest sites. A marked rise in temperature was recorded in nests from 30% to 40% of the way through the incubation period and attributed to metabolic heating. Over the entire incubation period, metabolic heating accounted for a mean rise in temperature of between 0.07 degrees and 2.86 degrees C within nests. During the middle third of incubation, when sex is thought to be determined, this rise in temperature ranged between 0.07 degrees and 2.61 degrees C. Metabolic heating was related to both the number of eggs laid and the total number of hatchlings/embryos produced in a clutch. For 32 clutches in which temperature was recorded, we estimate that metabolic heating accounted for a rise of up to 30% in the proportion of females produced within different clutches. Previous studies have dismissed any effect of metabolic heating on the sex ratio of marine turtle hatchlings. Our results imply that metabolic heating needs to be considered when estimating green turtle hatchling sex ratios.

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.

Open-sea migration of magnetically disturbed sea turtles.

Green turtles (Chelonia mydas) that shuttle between their Brazilian feeding grounds and nesting beaches at Ascension Island in the middle of the Atlantic Ocean are a paradigmatic case of long-distance oceanic migrants. It has been suggested that they calculate their position and the direction of their target areas by using the inclination and intensity of the earth's magnetic field. To test this hypothesis, we tracked, by satellite, green turtles during their postnesting migration from Ascension Island to the Brazilian coast more than 2000 km away. Seven turtles were each fitted with six powerful static magnets attached in such a way as to produce variable artificial fields around the turtle that made reliance on a geomagnetic map impossible. The reconstructed courses were very similar to those of eight turtles without magnets that were tracked over the same period and in the previous year, and no differences between magnetically disrupted and untreated turtles were found as regards navigational performance and course straightness. These findings show that magnetic cues are not essential to turtles making the return trip to the Brazilian coast. The navigational mechanisms used by these turtles remain enigmatic.

The implications of variable remigration intervals for the assessment of population size in marine turtles.

Sea turtles nest on sandy beaches and tend to show high fidelity to specific nesting areas, but, despite this fidelity, the inter-annual variation in nesting numbers may be large. This variation may reflect the fact that turtles do not usually nest in consecutive years. Here, theoretical models are developed in which the interval between successive nesting years (the remigration interval) reflects conditions encountered on the feeding grounds, with good feeding years leading to a reduction in the remigration interval and vice versa. These simple models produce high levels of inter-annual variation in nesting numbers with, on occasion, almost no turtles nesting in some years even when the population is large and stable. The implications for assessing the size of sea turtle populations are considered.

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 1777-2342 km in 33-47 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.

The functional significance of ventilation frequency, and its relationship to oxygen demand in the resting brown long-eared bat, Plecotus auritus.

Mean oxygen consumption and simultaneous ventilation frequency of nine non-reproductive brown long-eared bats (body mass 8.53-13.33 g) were measured on 159 occasions. Ambient (chamber) temperature at which the measurements were made ranged from 10.8 to 41.1 degrees C. Apneic ventilation occurred in 22 of the 59 measurements made when mean oxygen consumption was less than 0.5 ml.min-1. No records of apneic ventilation were obtained when it was over 0.5 ml.min-1. The relationship between ventilation frequency and mean oxygen consumption depended on whether ventilation was apneic or non-apneic. When ventilation was non-apneic the relationship was positive and log-linear. When ventilation was apneic the relationship was log-log. Within the thermoneutral zone ventilation frequency was not significantly different from that predicted from allometric equations for a terrestrial mammal of equivalent body mass, but was significantly greater than that predicted for a bird. A reduction in the amount of oxygen consumed per breath occurred at ambient temperatures above the upper critical temperature (39 degrees C).

Arrhythmic breathing in torpid pipistrelle bats, Pipistrellus pipistrellus.

The arrhythmic breathing pattern of torpid female pipistrelle bats (Pipistrellus pipistrellus) was monitored using Doppler radar. A total of 98 h of radar measurements were made on 11 individuals over 17 experiments, during which time 974 apneic intervals were monitored, over ambient temperatures (Ta, degrees C) ranging from -1 to 14 degrees C, and body masses ranging from 4.6 to 7.4 g. As Ta declined, a greater proportion of all breaths occurred in discrete breathing bouts. Apneic intervals lengthened at lower Ta, but were not related to body mass. Mean apneic length, averaged over 1 degree C intervals, was best described by the least squares fit regression equation: ln (apneic length in s) = 7.07-0.811 ln (Ta + 1), (r2 = 0.96, P less than 0.01). Ventilation frequency (breaths.min-1), averaged over a breathing bout and the subsequent apnea, increased as Ta increased, and was not related to body mass. Mean ventilation frequency (f), averaged over 1 degree C intervals, was best described by the least squares fit regression equation: f = 0.812 + 0.499 Ta (r2 = 0.92, P less than 0.01). Using previously published values for O2 consumption (VO2) in torpid pipistrelles, and tidal volume and O2 extraction efficiency at 4 degrees C in torpid bats of the same mean size (6.2 g), we calculated that at 4 degrees C ventilation would, on average, supply only 14.2-21.3% of VO2. This suggests that in torpid pipistrelles the glottis may remain open during apnea, allowing a significant diffusive influx of O2 into the lungs.