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.

Global research priorities for sea turtles : informing management and conservation in the 21st century

Over the past 3 decades, the status of sea turtles and the need for their protection to aid population recovery have increasingly captured the interest of government agencies, non-governmental organisations (NGOs) and the general public worldwide. This interest has been matched by increased research attention, focusing on a wide variety of topics relating to sea turtle biology and ecology, together with the interrelations of sea turtles with the physical and natural environments. Although sea turtles have been better studied than most other marine fauna, management actions and their evaluation are often hindered by the lack of data on turtle biology, human–turtle interactions, turtle population status and threats. In an effort to inform effective sea turtle conservation a list of priority research questions was assembled based on the opinions of 35 sea turtle researchers from 13 nations working in fields related to turtle biology and/or conservation. The combined experience of the contributing researchers spanned the globe as well as many relevant disciplines involved in conservation research. An initial list of more than 200 questions gathered from respondents was condensed into 20 metaquestions and classified under 5 categories: reproductive biology, biogeography, population ecology, threats and conservation strategies.

Breeding periodicity for male sea turtles, operational sex ratios, and implications in the face of climate change

Species that have temperature-dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long-standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single-sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in-water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female-biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.

Microhabitat selection by sea turtles in a dynamic thermal marine environment

1Reproductive fitness is often compromised at the margins of a species’ range due to sub-optimal conditions.2Set against this backdrop, the Mediterranean's largest loggerhead sea turtle (Caretta caretta) rookery at Zakynthos (Greece) presents a conundrum, being at a very high latitude for this species, yet hosting a high concentration of nesting.3We used visual surveys combined with global positioning system (GPS) tracking to show that at the start of the breeding season, individuals showed microhabitat selection, with females residing in transient patches of warm water. As the sea warmed in the summer, this selection was no longer evident.4As loggerhead turtles are ectothermic, this early season warm-water selection presumably speeds up egg maturation rates before oviposition, thereby allowing more clutches to be incubated when sand conditions are optimal during the summer.5Active selection of warm waters may allow turtles to initiate nesting at an earlier date.

Overwintering behaviour in sea turtles : dormancy is optional

Thirteen loggerhead turtles Caretta caretta were released (10 from Naples, Italy, 2 from Monastir, Tunisia, 1 from Gallipoli, South Italy) with satellite relay data loggers (SRDL) to elucidate their overwintering behaviour. Nine turtles were successfully tracked throughout the winter, while 4 SRDLs failed to transmit after short deployment periods. Of these 9, 4 remained within 80 km of the release site, 3 travelled to a distant overwintering site, and 2 continued to move and did not remain within 80 km of a specific site. Apart from these differences, all turtles stayed near the coast and dedicated most of their time to dives lasting 3 h and longer. Maximum dive durations ranged from 270 to 480 min and were highly correlated with water temperatures, which fell below the supposed 15°C threshold for sea turtle hibernation in all overwintering sites. Median dive depths were between 4 and 24 m and were, thus, well within the mixed layer, as revealed by temperature profiles, which also were relayed by the SRDLs. No evidence was found that the turtles preferred warmer temperatures to overwinter in, because the range of temperature was very narrow on both the horizontal and the vertical scale of their movements. Despite the long resting phases and the low temperatures (minimum = 11.8°C) all turtles retained activity to some degree, at least to commute between the depth of resting and the surface to breathe. While the degree of winter dormancy is certainly affected by temperature, turtles were by no means obligatory hibernators, and their ability to move and even forage during the winter may be important for their growth and maturation rates, as well as their reproductive output.

Female–female aggression : structure of interaction and outcome in loggerhead sea turtles

Aggressive behaviour between females of the same species is not widely documented, particularly in marine vertebrates. During a 3 yr in-water survey at the temperate loggerhead sea turtle Caretta caretta breeding area of Zakynthos, Greece, female–female interactions comprised 4% of all female loggerhead sighting events (n = 60 out of 1449 events). Male–female interactions comprised an additional 4% of sighting events, while 92% were of solitary females. The structure of interactions was analysed for 58 of these sighting events, each lasting an average of 3.4 min (SD ± 1) and comprising a total of 3.1 h observation time. We found that interactions involved ritualized escalation in behaviour from passive threat displays (e.g. head–tail circling) to aggressive combat (e.g. sparring). We suggest that circling individuals evaluate opponent size, sparring individuals test opponent strength, and that the positioning of the prehensile tail signals motivational intent to either escalate or abort. The presence of intruder females triggered a passive response in 100% of events involving basking and swimming turtles (n = 19); although residents resting on the seabed only responded on 69% of occasions (n = 27), their response was almost 4 times more likely to escalate to one of aggression. Our results suggest that certain sites may be preferentially sought after and defended by sea turtles.

Different male vs. female breeding periodicity helps mitigate offspring sex ratio skews in sea turtles

 The implications of climate change for global biodiversity may be profound with those species with little capacity for adaptation being thought to be particularly vulnerable to warming. A classic case of groups for concern are those animals exhibiting temperature-dependent sex-determination (TSD), such as sea turtles, where climate warming may produce single sex populations and hence extinction. We show that, globally, female biased hatchling sex ratios dominate sea turtle populations (exceeding 3:1 in >50% records), which, at-a-glance, reiterates concerns for extinction. However, we also demonstrate that more frequent breeding by males, empirically shown by satellite tracking 23 individuals and supported by a generalized bio-energetic life history model, generates more balanced operational sex ratios (OSRs). Hence, concerns of increasingly skewed hatchling sex ratios and reduced population viability are less acute than previously thought for sea turtles. In fact, in some scenarios skewed hatchling sex ratios in groups with TSD may be adaptive to ensure optimum OSRs.

Predicting climate warming effects on green turtle hatchling viability and dispersal performance

Ectotherms are taxa considered highly sensitive to rapid climate warming. This is because body temperature profoundly governs their performance, fitness and life history. Yet, while several modelling approaches currently predict thermal effects on some aspects of life history and demography, they do not consider how temperature simultaneously affects developmental success and offspring phenotypic performance, two additional key attributes that are needed to comprehensively understand species responses to climate warming. Here, we developed a stepwise, individual-level modelling approach linking biophysical and developmental models with empirically derived performance functions to predict the effects of temperature-induced changes to offspring viability, phenotype and performance, using green sea turtle hatchlings as an ectotherm model. Climate warming is expected to particularly threaten sea turtles, as their life-history traits may preclude them from rapid adaptation. Under conservative and extreme warming, our model predicted large effects on performance attributes key to dispersal, as well as a reduction in offspring viability. Forecast sand temperatures produced smaller, weaker hatchlings, which were up to 40% slower than at present, albeit with increased energy stores. Conversely, increases in sea surface temperatures aided swimming performance. Our exploratory study points to the need for further development of integrative individual-based modelling frameworks to better understand the complex outcomes of climate change for ectotherm species. Such advances could better serve ecologists to highlight the most vulnerable species and populations, encouraging prioritization of conservation effort to the most threatened systems.

Satellite tracking large numbers of individuals to infer population level dispersal and core areas for the protection of an endangered species

Aim: Tracking the dispersal patterns and habitat use of migratory species is necessary to delineate optimal areas for protection, with large sample sizes being more representative of the population. Here, we examine the dispersal patterns of a key Mediterranean loggerhead turtle (Caretta caretta) breeding population to identify priority foraging sites for protection. Location: Zakynthos Island, Greece and the wider Mediterranean. Method: We examined the dispersal patterns and foraging sites of 75 adult loggerheads (n = 38 males and 37 females) tracked from the breeding area of Zakynthos Island (Greece) from 2004 to 2011. We then combined our data with published sea turtle literature to identify key foraging sites for protection. Results: While both males and females exhibited similar dispersal patterns, about 25% males remained < 100 km of Zakynthos, whereas all females (except one) migrated > 200 km. Integration of our data with the wider literature isolated 10 core sites in proximity to existing protected areas, which could potentially protect 64% of the Zakynthos population, while five sites support individuals from at least 10 other loggerhead breeding populations. Main conclusions: Due to the widespread availability of neritic foraging grounds across the Mediterranean, sea turtles from Zakynthos exhibit disparate dispersal patterns. However, protecting only a few objectively defined important sites can encompass a large proportion of the foraging areas used and hence have considerable conservation benefit.

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.

Habitat utilization by juvenile hawksbill turtles (Eretmochelys imbricata, Linnaeus, 1766) around a shallow water coral reef

Contemporary studies of sea turtle diving behaviour are generally based upon sophisticated techniques such as the attachment of time depth recorders. However, if the risks of misinterpretation are to be minimized, it is essential that electronic data are analysed in the light of first-hand observations. To this aim, we set out to make observations of juvenile hawksbill turtles (Eretmochelys imbricata, Linnaeus, 1766) foraging and resting in a shallow water coral reef habitat around the granitic Seychelles (4°'S, 55°'E). Data were collected from six study sites characterized by a shallow reef plateau (<5 m) and a flat sandy area at the base of the reef face (<10 m). Observation data were categorized into the following behaviours: (1) stationary foraging; (2) active foraging; (3) resting; and (4) assisted resting. Central to this investigation was the development of a technique for accurately estimating the size of sea turtles in situ based upon previously tested techniques for reef fishes. This revealed that through calibration, the curved carapace length (CCL) of marine turtles can be consistently estimated to within 10 cm of their actual size. Although rudimentary, this has advantages for assessing the residency or absence of specific life history stages from particular environments. Indeed, our data supported previous claims that following the reproductive season, adult hawksbills in the region may move away from the nesting beaches to alternative foraging grounds whilst immature turtles (following the pelagic juvenile stage) may opt to reside in areas close to their natal beaches. With regards to habitat utilization, juvenile hawksbills displayed an alternating pattern of short, shallow foraging dives followed by deeper, longer resting dives. These findings are consistent with previous electronic studies of free-range diving in this species and suggest that the maximization of resting duration may be an important factor driving this behaviour.