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.

Thermal conductivity of sand and its effect on the temperature of Loggerhead Sea Turtle (Caretta Caretta) nests

The conductivity of sand at a depth of 30–50 cm was measured at 15 sites on the beach at Captiva Island in south-west Florida which is used by nesting loggerhead turtles (Caretta caretta). The mean daily temperature of the sand was correlated with conductivity at the same depth measured the same day (r=0·611). When day to day variation was removed the correlation between nest temperature and conductivity increased to 0·694. The sand was highly variable in its grain structure. The dominant variability (80·6%) was redescribed by the first two principal components of a Principal Components Analysis (PCA). These two components were influenced mostly by percentages of large (> 1 mm) and small (< 500 μm) grains respectively. Conductivity was strongly correlated with the grain structure of the sand. The first three principal components describing sand grain structure, explained 84·1% of the variation in conductivity. Moisture content of the sand (always < 5%) was not an important factor. Sites dominated by larger grains generally had poorer conductivity and were cooler. Comparisons of eight nests to seven adjacent random sites revealed no strong evidence for directional selection in nest placement relative to sand conductivity. The variance in conductivities recorded at nests was also not significantly different from the variance at random sites.

First records of dive durations for a hibernating sea turtle

The first published record, from the early 1970s, of hibernation in sea turtles is based on the reports of the indigenous Indians and fishermen from Mexico, who hunted dormant green turtles (Chelonia mydas) in the Gulf of California. However, there were no successful attempts to investigate the biology of this particular behaviour further. Hence, data such as the exact duration and energetic requirements of dormant winter submergences are lacking. We used new satellite relay data loggers to obtain the first records of up to 7 h long dives of a loggerhead turtle (Caretta caretta) overwintering in Greek waters. These represent the longest dives ever reported for a diving marine vertebrate. There is strong evidence that the dives were aerobic, because the turtle surfaced only for short intervals and before the calculated oxygen stores were depleted. This evidence suggests that the common belief that sea turtles hibernate underwater, as some freshwater turtles do, is incorrect.

Employing sea-level rise scenarios to strategically select sea turtle nesting habitat important for long-term management at a temperate breeding area

Management strategies to protect endangered species primarily focus on safeguarding habitats currently perceived as important (due to high-density use, rarity or contribution to the biological cycle), rather than sites of future ecological importance. This discrepancy is particularly relevant for species inhabiting beaches and coastal areas that may be lost due to sea-level rise over the next 100 years through climate change. Here, we modelled four sea-level rise (SLR) scenarios (0.2, 0.6, 0.9 and 1.3 m) to determine the future vulnerability and viability of nesting habitat (six distinct nesting beaches totalling about 6 km in length) at a key loggerhead sea turtle (Caretta caretta) rookery (Zakynthos, Greece) in the Mediterranean. For each of the six nesting beaches, we identified (1) the area of beach currently used by turtles, (2) the area of the beach anticipated to become inundated under each SLR, (3) the area of beach anticipated to become unsuitable for nesting under each SLR, (4) the potential for habitat loss under the examined SLR, and (5) the extent to which the beaches may shift in relation to natural (i.e. cliffs) and artificial (i.e. beach front development) physical barriers. Even under the most conservative 0.2 m SLR scenario, about 38% (range: 31–48%) total nesting beach area would be lost, while an average 13% (range: 7–17%) current nesting beach area would be lost. About 4 km length of nesting habitat (representing 85% of nesting activity) would be lost under the 0.9 m scenario, because cliffs prevent landward beach migration. In comparison, while the other 2 km of beach (representing 15% nests) is also at high risk, it has the capacity for landward migration, because of an adjoining sand-dune system. Therefore, managers should strengthen actions on this latter area, as a climatically critical safeguard for future sea turtle nesting activity, in parallel to regularly assessing and revising measures on the current high-use nesting habitats of this important Mediterranean loggerhead population.

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.

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.

Acceleration data reveal the energy management strategy of a marine ectotherm during reproduction

1. Maintaining a high and stable body temperature is often critical for female ectotherms during reproduction. Yet this strategy may be energetically costly, and therefore challenging, during this period of already high-energy demand. 2. Here, the 6-week deployment of tri-axial accelerometers (n = 6) on a marine ectotherm, the loggerhead turtle (Caretta caretta), reproducing at the northern limit of the species’ breeding range (i.e. in a thermally dynamic environment) revealed the behavioural mechanisms underlying its energy management strategy during the breeding season. 3. The estimated activity levels of female loggerheads using overall dynamic body acceleration (ODBA) were high during the breeding season, suggesting that marine turtles may not be able to remain inactive for long periods in the same manner as terrestrial ectotherms, because of the thermally dynamic nature of their environment. 4. However, activity levels were not constant throughout the season, being impacted by both ambient water temperature and female reproductive status. In cold water at the beginning of the nesting season, high levels of activity suggested that females behaviourally thermoregulated by seeking out warm water patches along the shoreline. Interactions with male turtles (courtship and/or avoidance) may also explain this high level of activity. As sea temperatures warmed up and the amount of energy devoted to reproduction probably increased, the turtles spent more time resting during long sequential flat-bottomed dives, and reduced any unnecessary locomotory activity. 5. Turtles may therefore adjust their activity patterns in response to seasonal variations in abiotic (i.e. ambient temperature) and biotic (i.e. reproductive status) factors. This may help minimize activity-linked metabolic rate and maximize reproductive output over a season while breeding in thermally dynamic environments. 6. A mechanistic model gave support to these empirical results. The model revealed that actively maintaining high and stable body temperature is of clear benefit to female turtles at temperate breeding sites. While energetically costly, such active thermoregulatory behaviour may speed up egg maturation, allowing turtles to initiate nesting earlier in the season, and hence maximize 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.

Living on the edge: how philopatry maintains adaptive potential.

Without genetic variation, species cannot cope with changing environments, and evolution does not proceed. In endangered species, adaptive potential may be eroded by decreased population sizes and processes that further reduce gene flow such as philopatry and local adaptations. Here, we focused on the philopatric and endangered loggerhead sea turtle (Caretta caretta) nesting in Cape Verde as a model system to investigate the link between adaptive potential and philopatry. We produced a dataset of three complementary genomic regions to investigate female philopatric behaviour (mitochondrial DNA), male-mediated gene flow (microsatellites) and adaptive potential (major histocompatibility complex, MHC). Results revealed genetically distinct nesting colonies, indicating remarkably small-scale philopatric behaviour of females. Furthermore, these colonies also harboured local pools of MHC alleles, especially at the margins of the population's distribution, which are therefore important reserves of additional diversity for the population. Meanwhile, directional male-mediated gene flow from the margins of distribution sustains the adaptive potential for the entire rookery. We therefore present the first evidence for a positive association between philopatry and locally adapted genomic regions. Contrary to expectation, we propose that philopatry conserves a high adaptive potential at the margins of a distribution, while asymmetric gene flow maintains genetic connectivity with the rest of the population.

Quantifying wildlife-watching ecotourism intensity on an endangered marine vertebrate

Here, we show how seasonal changes in animal density drive strategic shifts in the activities of wildlife-watching operators. These shifts result in high viewing intensity when animal densities are low, highlighting the need for modifications to existing wildlife-watching guidelines. We used the endangered loggerhead sea turtle Caretta caretta as a model species that exhibits staggered departure from an important breeding area (Zakynthos, Greece, Mediterranean) over a 2-month period (July to August) when tourism is at a peak, to investigate changes in wildlife-watching strategies, zoning effectiveness and voluntary guideline compliance over time. We used a combination of direct land-based observations, global positioning system tracking (of wildlife-watching vessels and turtles) and models. The modelled number of turtles present in the breeding area decreased from >200 in July to <50 in August, while the intensity of turtle-viewing increased from a mean 1.5 to 6.1 wildlife-watching vessels per turtle-viewing event (i.e. concurrent and consecutive vessels observing a single turtle) over the same period, respectively. During this period, the wildlife-watching strategy changed and compliance to guidelines reduced (exacerbated by recreational vessels). However, wildlife-watching activity was limited to a highly restricted 0.95-km2 nearshore area, overlapping with just 9.5% of the core habitat area used by turtles. Our results have broad implications (whale watching etc.) by showing the importance of taking the number of animals available for viewing into consideration when assessing wildlife-watching activity and when designing viewing guidelines, particularly for populations where numbers noticeably fluctuate.

Patterns in the emergence of green (Chelonia mydas) and loggerhead (Caretta caretta) turtle hatchlings from their nests

The emergence patterns of both green (Chelonia mydas) and loggerhead (Caretta caretta) turtle hatchlings were observed in great detail over three seasons at Alagadi beach, northern Cyprus. In total, 38 green turtle and 50 loggerhead turtle nests were monitored, accounting for the emergence of 2,807 and 2,259 hatchlings, respectively. We quantified these emergences into 397 green turtle and 302 loggerhead turtle emergence groups. Overall, 85.0% of green turtle and 79.5% of loggerhead turtle groups emerged at night; these accounted for 85.5 and 90.8% of hatchlings, respectively. The remaining emergences were dispersed throughout the day for green turtle nests but confined to the morning in loggerhead turtle nests. Hatchling emergence from individual nests occurred over periods of between 1 and 7 nights, with most hatchlings typically emerging on the first night. Group sizes of green turtles emerging during the day were significantly smaller than those emerging at night. Hatchlings of both species that emerged from nests during the day had longer emergence durations than those that emerged from nests at night only.

Diving behaviour during the internesting interval for loggerhead turtles Caretta caretta nesting in Cyprus

Time depth recorders were used to assess the patterns of depth utilisation by 2 loggerhead turtles Caretta caretta in Cyprus, eastern Mediterranean. Dives to the seabed accounted for 59% (171 h) and 75% (215 h) of the internesting interval, respectively, with most dives being shallow (<20 m), suggesting the turtles remained close to the shore. These benthic dives decreased markedly in the days following or prior to a nesting event, suggesting that the behaviours associated with nesting may be protracted. This importance of the seabed for loggerhead turtles in Cyprus contrasts with the far more extensive use of mid-water resting dives recently reported for this species in Japan. Our evidence suggests that this dichotomy may reflect differences in the amount of time spent travelling, with mid-water resting occurring when turtles are travelling and, conversely, when little time is spent travelling turtles opt to remain predominantly on the seabed.

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.

Two hundred years after a commercial marine turtle fishery: the current status of marine turtles nesting in the Cayman Islands

Large populations of marine turtles breeding in the Cayman Islands were drastically reduced in the early 1800s. However, marine turtle nesting still occurs in the islands. The present-day status of this nesting population provides insight into the conservation of marine turtles, a long-lived species. In 1998 and 1999, the first systematic survey of marine turtle nesting in the Cayman Islands found 38 nests on 22 beaches scattered through the three islands. Three species were found: the green Chelonia mydas, hawksbill Eretmochelys imbricata and loggerhead Caretta caretta turtles. Comparison with other rookeries suggests that the small number of sexually mature adults surviving Cayman’s huge perturbations may be impeding population recovery. This shows the need to implement conservation measures prior to massive reductions in population size.