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The loggerhead sea turtle (Caretta caretta), has the widest distribution among sea turtle species in the Mediterranean, requiring regional and international collaborations in addition to local efforts to better inform conservation actions. Molecular techniques are powerful tools for the assessment of population dynamics at large scales, especially in determining the connectivity among different nesting and foraging sites, and genetic diversity. In this study, a large sample was collected synchronously in the nesting areas located in the north, south and eastern Mediterranean. Recently confirmed nesting sites from Albania and other nesting sites represented by lower sample sizes were also included in order to better assess the genetic composition of the region's rookeries. Samples from 698 individuals were collected and the longer (815 bp) mtDNA D-loop fragment of these samples was sequenced. We recorded 15 haplotypes, three of which were novel. In addition, our results show that some haplotypes considered of Atlantic origin, have a wider dispersal in the Mediterranean than previously thought, albeit with low levels of representation. Our results contribute to determining the likely origin of haplotypes that were previously only recorded from foraging sites. They highlight the utility of broad-scale sampling, with increased sample sizes and the longer mtDNA sequence to determine genetic diversity and connectivity. This study also demonstrates the importance of continued monitoring for the contribution of Atlantic-origin haplotypes to the Mediterranean population and the effects of climate change on the resident Mediterranean population, which is expected to expand its geographical range for reproduction. This work is important for, mixed stock analyses (MSA) that seek to determine the natal regions of stranded or accidentally caught sea turtles and those purposefully obtained from foraging sites. In doing so biogeographic linkages between areas of the Mediterranean can be elucidated for conservation purposes.

Assessing sea turtle movements and connectivity among different areas is pivotal to understanding their biology and implementing efficient conservation actions. In the Adriatic Sea, one of the most important sea turtle foraging areas in the Mediterranean, a total of 311 capture–mark–recapture (CMR) records (mostly bycatch) from 294 loggerhead turtles (Caretta caretta) in the period 1984–2021 were analysed. A general fidelity pattern to Adriatic subareas was indicated by a significantly shorter CMR distance than the potential dispersal distance and by the significantly higher proportion of re-encounters in the same area of release than expected. No seasonal pattern was detected between subareas, and shorter re-encounter distances were observed in turtles released and re-encountered in the same season, suggesting different winter and summer residential areas. Results suggest that turtles frequenting the Adriatic can go anywhere in the Mediterranean basin and may exhibit a wandering behaviour regardless of their size. A substantial connectivity with nesting sites in Greece was observed, confirming with empirical evidence that this is the most important breeding area for turtles foraging in the Adriatic Sea. This study highlights the value of cooperation among different groups and shows a main behavioural pattern of fidelity to neritic foraging grounds.

Aim: Understanding the spatial ecology of animal movements is a critical element in conserving long-lived, highly mobile marine species. Analyzing networks developed from movements of six sea turtle species reveals marine connectivity and can help prioritize conservation efforts. Location: Global. Methods: We collated telemetry data from 1235 individuals and reviewed the literature to determine our dataset's representativeness. We used the telemetry data to develop spatial networks at different scales to examine areas, connections, and their geographic arrangement. We used graph theory metrics to compare networks across regions and species and to identify the role of important areas and connections. Results: Relevant literature and citations for data used in this study had very little overlap. Network analysis showed that sampling effort influenced network structure, and the arrangement of areas and connections for most networks was complex. However, important areas and connections identified by graph theory metrics can be different than areas of high data density. For the global network, marine regions in the Mediterranean had high closeness, while links with high betweenness among marine regions in the South Atlantic were critical for maintaining connectivity. Comparisons among species-specific networks showed that functional connectivity was related to movement ecology, resulting in networks composed of different areas and links. Main conclusions: Network analysis identified the structure and functional connectivity of the sea turtles in our sample at multiple scales. These network characteristics could help guide the coordination of management strategies for wide-ranging animals throughout their geographic extent. Most networks had complex structures that can contribute to greater robustness but may be more difficult to manage changes when compared to simpler forms. Area-based conservation measures would benefit sea turtle populations when directed toward areas with high closeness dominating network function. Promoting seascape connectivity of links with high betweenness would decrease network vulnerability. © 2022 The Authors. Diversity and Distributions published by John Wiley & Sons Ltd.

In sea turtles, somatic growth rates and Age at Sexual Maturity (ASM) are important parameters for investigating population dynamics. Moreover, ASM informs on the time lag needed to observe the effects of past environmental variables, threats and conservation measures acting at nesting beaches on future numbers of clutches or nesting females, that are the most monitored indices. Assessing ASM of mixed populations that share the same foraging areas is difficult, as in the case of the Mediterranean Sea that is frequented by turtles originating from both the Mediterranean and the Atlantic. Here we investigated growth rates of loggerhead sea turtles (Caretta caretta) foraging in the Adriatic Sea, known to be frequented only by individuals of Mediterranean origin. Capture–mark–recapture records of 79 turtles provided growth rates ranging from – 1.1 to 9.6 cm yr−1 which were analyzed through Generalized Additive Models. When integrated with previous data on growth rates in the first years of life, results indicate a non-monotonic growth curve, likely due to an ontogenetic shift from oceanic to neritic areas. Using an integration equation, we estimated mean ASM to be 29.5 or 25.0 year based on mean nester size in Greece (83.4 cm curved carapace length, CCL) or across the Mediterranean (79.7 cm CCL), respectively. Growth rates and size-at-age are similar to the Atlantic population, but Mediterranean turtles mature earlier at a smaller size. The heterogenous growth rates and ASM reported so far in the Mediterranean are probably due, at least in part, to different foraging areas.

The loggerhead sea turtle (Caretta caretta), has the broadest distribution among sea turtle species in the Mediterranean and requires regional and international collaborations in addition to local efforts to better inform conservation actions. Molecular techniques are powerful tools to assess population dynamics at large scales, especially by determining the connectivity among different nesting and foraging sites, and genetic diversity. In this study, a large sample was collected synchronously in the nesting areas located in the north, south and east of the Mediterranean. Recently described nesting sites from Albania and other nesting sites represented by lower sample size were also included in order to fully assess the genetic composition of the region’s rookeries. Samples from 710 individuals were collected and the longer (815 bp) mtDNA D-loop fragment of these samples was amplified. We recorded 15 haplotypes, three of which were novel. In addition, our results show that some haplotypes, considered of Atlantic origin, have a wider dispersal in the Mediterranean than previously thought, albeit with low levels of representation. Our results, which also contribute to determining the likely origin of haplotypes that were previously known only from foraging sites, highlight the utility of broad-scale sampling, with increased sample number and longer mtDNA sequence to determine genetic diversity and connectivity. This study also demonstrates that it is important to continue to monitor the contribution of Atlantic origin haplotypes to the Mediterranean population, and the resident Mediterranean population, which is expected to expand its geographical range for reproduction with the effect of climate change and climate change in the long term. This work is important for, among other things, mixed stock analyses (MSA) that seek to localize the origin of stranded or accidentally caught sea turtles or those purposefully obtained from foraging sites to better understand the migratory distribution for conservation purposes.

AIM: Using a combination of satellite telemetry and stable isotope analysis (SIA), our aim was to identify foraging grounds of loggerhead turtles (Caretta caretta) at important rookeries in the Mediterranean, examine foraging ground fidelity, and across 25 years determine the proportion of nesting females recruiting from each foraging region to a major rookery in Cyprus. LOCATION: Mediterranean Sea. METHODS: Between 1993 and 2018, we investigated the spatial ecology of loggerhead turtles from rookeries in Cyprus and Greece using satellite telemetry (n = 55 adults) and SIA of three elements (n = 296). RESULTS: Satellite telemetry from both rookeries revealed the main foraging areas as the Adriatic region (Cyprus: 4% of individuals, Greece: 55%), Tunisian Plateau (Cyprus: 16%, Greece: 40%) and the eastern Mediterranean (Cyprus: 80%, Greece: 5%). Combining satellite telemetry and SIA allowed 64% of all nesting females to be assigned to; the Adriatic region (Cyprus: 2%, Greece: 38.5%), Tunisian Plateau (Cyprus: 47%, Greece: 38.5%) and the eastern Mediterranean (Cyprus: 51%, Greece: 23%), which are markedly different to proportions obtained using satellite telemetry. The proportion of the Cyprus nesting cohort using each foraging region did not change significantly, with the exception that individuals foraging in the Adriatic region are only present in the Cyprus nesting population from 2012. Repeat satellite tracking (n = 3) and temporal consistency in isotope ratios (n = 36) of Cyprus females, strongly suggest foraging ground fidelity over multiple decades. MAIN CONCLUSIONS: This study demonstrates the advantages of combining satellite telemetry and SIA to investigate spatial ecology at a population level. The importance of the Tunisian Plateau for foraging is demonstrated. This study indicates that females generally show high fidelity to foraging grounds and shows a potential recent shift to foraging in the Adriatic region for Cyprus females, while the importance of other regions persists across decades, thus providing baselines to develop and assess conservation strategies.

Bycatch is one of the key threats to juvenile marine turtles in the Mediterranean Sea. As fishing methods are regional or habitat specific, the susceptibility of marine turtles may differ according to inter- and intra-population variations in foraging ecology. An understanding of these variations is necessary to assess bycatch susceptibility and to implement region-specific management. To determine if foraging ecology differs with region, sex, and size of juvenile loggerhead turtles (Caretta caretta), stable isotope analysis of carbon and nitrogen was performed on 171 juveniles from a range of foraging regions across the central and eastern Mediterranean Sea. Isotope ratios differed with geographical region, likely due to baseline variations in δ13C and δ15N values. The absence of sex-specific differences suggests that within an area, all comparably sized animals likely exploit similar foraging strategies, and therefore, their susceptibility to fisheries threats will likely be similar. The isotope ratios of juveniles occupying the North East Adriatic and North Levantine basin increased with size, potentially due to increased consumption of more prey items at higher trophic levels from a more neritic source. Isotope ratios of juveniles with access to both neritic and oceanic habitats did not differ with size which is consistent with them consuming prey items from both habitats interchangeably. With foraging habitats exploited differently among size classes in a population, the susceptibility to fisheries interactions will likely differ with size; therefore, region-specific management approaches will be needed.

Following a multi-decadal tagging program at loggerhead nesting areas in Greece, 28 individual turtles have been identified with reproductive longevities greater than 21 yrs, ranging from 22 to 33 yrs. Continuation of this long-term tagging program, along with improvement of tag retention through usage of PIT tags, are expected to increase the herein reported maximum reproductive longevity of loggerhead turtles.

The available information regarding the 2 sea turtle species breeding in the Mediterranean (loggerhead turtle Caretta caretta and green turtle Chelonia mydas) is reviewed, including biometrics and morphology, identification of breeding and foraging areas, ecology and behaviour, abundance and trends, population structure and dynamics, anthropogenic threats and conservation measures. Although a large body of knowledge has been generated, research efforts have been inconsistently allocated across geographic areas, species and topics. Significant gaps still exist, ranging from the most fundamental aspects, such as the distribution of major nesting sites and the total number of clutches laid annually in the region, to more specific topics like age at maturity, survival rates and behavioural ecology, especially for certain areas (e.g. south-eastern Mediterranean). These gaps are particularly marked for the green turtle. The recent positive trends of nest counts at some nesting sites may be the result of the cessation of past exploitation and decades of conservation measures on land, both in the form of national regulations and of continued active protection of clutches. Therefore, the current status should be considered as dependent on such ongoing conservation efforts. Mitigation of incidental catch in fisheries, the main anthropogenic threat at sea, is still in its infancy. From the analysis of the present status a comprehensive list of re search and conservation priorities is proposed., C.C. is supported by the project CTM2013-48163 of the Spanish Ministry of Economy and Competitivity. The Cyprus Wildlife Society (CWS) acknowledges the financial support of the Department of Fisheries and Marine Research of the Government to the CWS for the implementation of the Turtle Conservation Project in the period 2010−2016 and for all its assistance to the Society in previous years. J.T. is supported by project Prometeo II (2015-018) of the Generalitat Valenciana and projects MEDSEALITTER-INTERREG and INDICIT of the European Union.

Sea turtle populations worldwide suffer from reduced survival of immatures and adults due to fishery bycatch. Unfortunately, information about the whereabouts of turtles outside the breeding habitat is scarce in most areas, hampering the development of spatially explicit conservation plans. In the Mediterranean, recoveries of adult females flipper-tagged on nesting beaches suggest that the Adriatic Sea and Gulf of Gabes are important foraging areas for adults, but such information could be heavily biased (observing and reporting bias). In order to obtain unbiased data, we satellite-tracked seven loggerhead sea turtles after they completed nesting in the largest known Mediterranean rookery (Bay of Laganas, Zakynthos, Greece). Three females settled in the north Adriatic Sea, one in the south Adriatic Sea and two in the Gulf of Gabes area at the completion of their post-nesting migrations (one individual did not occupy a distinct foraging area). The concordance of tracking results with information from recoveries of flipper-tagged turtles suggests that the north Adriatic Sea and the Gulf of Gabes represent key areas for female adult Mediterranean loggerhead sea turtles.

Marine turtles have been traditionally considered model organisms to study sex-biased behaviour and dispersal. Although female philopatry has been identified in the loggerhead turtle, with adult females returning to specific locations to nest, studies on the philopatry and breeding migrations of males remain limited. In this study we analysed 152 hatchlings using 15 microsatellite markers. Each individual came from a different nest from samples taken at 8 nesting grounds in the Mediterranean. Our results revealed the existence of 5 genetically differentiated units, mostly due to restricted gene flow for both sexes. This supports existing satellite tracking studies that suggest that mating occurs close to nesting grounds in this region. The 5 management units identified within the Mediterranean included nesting grounds from (1) Libya and Cyprus, (2) Israel, (3) Lebanon, (4) Turkey and (5) Greece. The genetic similarity between distant nesting areas (i.e. Libya and Cyprus) suggests the presence of a more complex pattern of breeding behaviour. Three possible hypotheses, that remain to be tested in future studies, could explain this result: (1) mating might take place in common foraging grounds; (2) mating could occur en route while migrating to/from the breeding grounds; or (3) recent colonisation events could connect the 2 nesting grounds. Overall, our work suggests that widespread male-mediated gene flow between loggerhead nesting grounds is likely to have been previously overstated although opportunistic breeding patterns might connect some widely separated areas.

Foraging success affects reproductive output in sea turtles, and is therefore an impor- tant factor to measure in order to understand population dynamics. During 2010 and 2011, we used satellite telemetry to track the at-sea behavior of 20 post-nesting loggerhead turtles Caretta caretta from Rethymno, Crete, Greece. Nineteen transmitters provided location and dive data throughout the turtles' migration towards their foraging grounds and the transition into foraging behavior. We identified 3 foraging regions: (1) 9 turtles migrated southwest towards the North African coast, with 8 concentrated in the region of the Gulf of Gabes, Tunisia; (2) 6 turtles migrated north towards the Aegean Sea; and (3) 4 turtles did not take long-distance migrations, instead remaining resident within the waters of Crete. Two fitness proxies were associated with differ- ences in post-nesting strategies. Turtles foraging in northern waters had significantly larger curved and straight carapace lengths and clutch sizes than turtles foraging near Crete or Africa. Those differences reflect the disparity in benthic prey abundances among the 3 regions. The Aegean had a higher abundance of macrobenthic fauna than the other 2 regions, and the Gulf of Gabes had an increased level of eutrophication. Deterioration of the aquatic resources in the Gulf of Gabes region may be a contributing factor in the observed steady decline in clutch size and total nests per season in 2 critical nesting beaches for loggerheads in Greece.

Small-scale fisheries are responsible for high numbers of animals caught as bycatch, such as turtles, cetaceans, and seals. Bycatch and its associated mortality is a major conservation challenge for these species and is considered undesirable by fishermen. To gain insights on the impact of bycatch on small-scale fishermen and put it in context with other financial and environmental challenges they face, we conducted questionnaire-based interviews on fishermen working on Crete, Greece. We investigated fishermen's perceptions of sea turtle and other protected species interactions, and the impacts of such interactions on their profession and livelihoods. Our results indicate a connection between declining fish stocks, related increased fishing effort, and reported increased frequency of interactions between fishermen and sea turtles. Respondents believed that their livelihoods were endangered by industrial fishing and environmental problems, but thought that combined interactions with turtles and other marine megafauna species were a larger problem. Responses suggested that extending compensation to fishermen may be a good conservation intervention. Small-scale fishermen hold a wealth of knowledge about the marine environment and its resources. This may be of help to researchers and policy makers as it could be used to achieve a better managed, sustainable fishery. Including small-scale fishermen in the process of developing regulations will both enhance those regulations and increase compliance with them.

Many marine megavertebrate taxa, including sea turtles, disperse widely from their hatching or birthing locations but display natal homing as adults. We used flipper tagging, satellite tracking and genetics to identify the origin of loggerhead turtles living in Amvrakikos Gulf, Greece. This location has been identified as hosting regionally important numbers of large-juvenile to adult sized turtles that display long-term residency and/or association to the area, and also presents a male biased sex ratio for adults. A total of 20 individuals were linked to nesting areas in Greece through flipper tagging and satellite telemetry, with the majority (16) associated with Zakynthos Island. One additional female was tracked from Amvrakikos Gulf to Turkey where she likely nested. Mitochondrial DNA mixed stock analyses of turtles captured in Amvrakikos Gulf (n = 95) indicated 82% of individuals originated from Greek nesting stocks, mainly from Zakynthos Island (63%), with lesser contributions from central Turkey, Cyprus and Libya. These results suggest that the male-biased sex ratio found in Amvrakikos Gulf may be driven by the fact that males breed twice as frequently on Zakynthos, resulting in their using foraging grounds of greater proximity to the breeding site. Conservation measures in localised foraging habitats for the protection of marine vertebrates, such as sea turtles, may have positive impacts on several disparate breeding stocks and the use of multiple methods to determine source populations can indicate the relative effectiveness of these measures. Electronic supplementary material The online version of this article (doi:10.1007/s00227-016-3055-z) contains supplementary material, which is available to authorized users.

Genetic markers have been widely used in marine turtles to assess population structuring and origin of individuals in common feeding grounds, which are key elements for understanding their ecology and for developing conservation strategies. However, these analyses are very sensitive to missing information, especially from abundant nesting sites. Kyparissia Bay (western Greece) hosts the second largest Mediterranean nesting aggregation of the loggerhead turtle (Caretta caretta), but the genetic profile of this nesting site has not, as yet, been described using the extended version of the historically used mitochondrial DNA (mtDNA) marker. This marker was genotyped for 36 individuals nesting at Kyparissia Bay and haplotype frequencies obtained were compared with published data from other Mediterranean nesting sites. The results confirmed the connection between Kyparissia and other western Greek nesting sites and the isolation of this western Greek group from other Mediterranean nesting areas. As a consequence of this isolation, this abundant group of nesting aggregations (almost 30% of the Mediterranean stock) is not likely to significantly contribute to the recovery of other declining Mediterranean units. El uso de marcadores genéticos ha sido muy extendido en tortugas marinas con el fin de determinar la estructura poblacional y el origen de individuos en zonas comunes de alimentación; elementos clave para entender su ecología y desarrollar estrategias efectivas de conservación. Sin embargo, este tipo de análisis es muy sensible a la falta de información de ciertas zonas de nidificación, especialmente de aquellas muy abundantes. El perfil genético de las tortugas nidificantes de la bahía de Kyparissia (Grecia occidental) todavía no ha sido descrito usando la versión extendida del marcador mitocondrial (mtDNA) usado históricamente en esta especie, a pesar de ser la segunda zona de nidificación más abundante de todo el Mediterráneo. Con el fin de cubrir este vacío de información, se secuenciaron 36 individuos nidificantes de la bahía de Kyparissia y se compararon las frecuencias de haplotipos obtenidas con datos publicados de otras zonas de nidificación del Mediterráneo. Los resultados confirmaron la conexión entre Kyparissia y otras zonas de nidificación de Grecia así como el aislamiento de este grupo de Grecia occidental con el resto de zonas de nidificación del Mediterráneo. Como consecuencia de este aislamiento, todo parece indicar que este abundante grupo de zonas de nidificación (casi el 30% de la producción del Mediterráneo) no podría contribuir de forma significativa a la recuperación del número de hembras nidificantes en otras poblaciones en declive del Mediterráneo.

Much is still to be learned about the spatial ecology of foraging marine turtles, especially for juveniles and adult males which have received comparatively little attention. Additionally, there is a paucity of ecological information on growth rates, size and age at maturity, and sex ratios at different life stages; data vital for successful population modelling. Here, we present results of a long-term (2002–2011) study on the movements, residency, growth and sex ratio of loggerhead turtles (Caretta caretta) in Amvrakikos Gulf (39°0′N 21°0′E), Greece, using satellite telemetry (N = 8) and ongoing capture–mark–recapture (CMR; N = 300 individuals). Individuals encountered at sea ranged from large juvenile to adult (46.2–91.5 cm straight carapace length) and demonstrated growth rates within published norms ( 44 % of captures above 65 cm straight carapace length), compared to region-wide female-biased hatchling production, indicating sex-biased survival or possible behavioural drivers for likelihood of capture in the region. Satellite tracking confirmed that some turtles establish discrete, protracted periods of residency spanning more than 1 year, whilst others migrated away from the site. These findings are underlined by CMR results with individual capture histories spanning up to 7 years, and only 18 % of individuals being recaptured.

Linking foraging and breeding habitats is key to the understanding of behaviour, ecol- ogy and demography of migratory species. Establishing such connections has long been hampered by the logistical problems of following individuals between foraging and breeding areas, especially in the marine realm. We used variation in nitrogen stable isotope patterns between 2 foraging regions of loggerhead sea turtles Caretta caretta determined from samples of satellite-tracked indi- viduals to assign untracked turtles to a foraging region. We sought to enhance determination of the relative importance of geographically separated foraging regions and to investigate the relationship between fitness correlates and inferred migratory strategies. Of 18 turtles followed by satellite tracking from Zakynthos (Greece), 10 moved north to foraging areas in the Adriatic Sea and the Gulf of Amvrakikos and 8 moved south to foraging areas off the coast of North Africa. Of 51 untracked individuals sampled for stable isotope analysis, we considered the stable isotope signature of 47 to qualify for assignment to foraging areas in the north (n = 22) and south (n = 25). Females foraging north were significantly larger (curved carapace length), and the former group laid larger clutches (even after correction for body length) than turtles foraging south, a fact that can be interpreted as a carry-over effect. Combining satellite tracking with stable isotope signatures in marine turtles opens new perspectives into how forensic tracking methodologies may be used to scale up knowl- edge from electronic tracking of a limited number of individuals to sample sizes that are more mean- ingful from a population perspective.

Young green turtles (Chelonia mydas) spend their early lives as oceanic omnivores, which consume primarily animal prey. Once they settle into neritic habitats (recruitment), they appear to shift rapidly into an herbivorous diet in tropical regions. However, the ontogeny of the dietary shift and the relevance of animal prey in the diet of neritic green turtles are poorly understood in subtropical and warm temperate regions. Stable isotopes of carbon and nitrogen in the carapace scutes of 22 green turtles from the eastern Mediterranean, ranging from 28 to 83 cm in curved carapace length (CCLmin), were analysed to test the hypothesis of a rapid nutritional shift following recruitment. Seagrasses prevailed in the stomach contents of all the green turtles that were larger than 30 cm CCLmin, but the concentration of stable isotopes in the carapace scutes revealed that turtles shorter than 40 cm CCLmin derived a negligible amount of assimilated nutrients from seagrasses. The concentration of stable isotopes in the carapace scutes also suggested that the contribution of seagrasses to the nutrients assimilated by green turtles increased steadily with turtle size and that some, but not all, of the turtles larger than 62 cm CCLmin were fully herbivorous. The overall evidence (gut contents analysis and stable isotope analysis) indicates that green turtles in the Mediterranean shift to a seagrass-based diet immediately after recruitment but turtle growth continues to rely on animal-derived nutrients for several years after recruitment. This asynchrony between the dietary and nutritional shifts is thought to be caused by the temperature sensitivity of bacterial fermentation and the low temperatures experienced by green turtles in the Mediterranean, which may result in a poor assimilation of the plant-derived nutrients for most of their neritic juvenile life.

The choice of a suitable nest habitat by oviparous reptiles that deposit eggs into a nest and provide no subsequent parental care is likely to play a major role in the survival of the offspring. In particular variations in nest temperature may influence the rate at which embryos utilise their yolk energy. The effects of nest temperature on total energy use are however complex. High temperatures may advance development and shorten the time to hatching, thereby reducing energy use, but they also stimulate metabolic rate increasing energy use. The net effect of temperature on total energy demands is therefore uncertain. Oxygen consumption (VO2) and carbon dioxide production (VCO2) were measured by open-flow respirometry during the incubation of loggerhead sea turtle eggs at three temperatures (27.6, 30.0 and 31.8 °C). At each temperature, VO2 and VCO2 showed a peak followed by a decline to hatching. Incubation temperature was negatively related to incubation duration and positively related to the maximum metabolic rate of the embryos. Peak VO2 was 74.8 ml/egg/day at 27.6 °C, 91.9 ml/egg/day at 30.0 °C, and 97.9 ml/egg/day at 31.8 °C. Peak VO2 occurred closer to hatching in eggs incubated at higher temperatures. Total energy expenditure was greatest at the lowest incubation temperature and lowest at the highest temperature. Total VO2 and VCO2 were 1777 ml/egg and 1226 ml/egg, respectively, at 27.6 °C, 1680 ml/egg and 1235 ml/egg at 30.0 °C, and 1613 ml/egg and 1191 ml/egg at 31.8 °C. Using the actual RQ values, this corresponds to a cost of development of 34,963 J/egg at 27.6 °C, 33,403 J/egg at 30.0 °C, and 32,107 J/egg at 31.8 °C. At all temperatures, the calculated respiratory quotient values did not suggest that yolk substrates were oxidised proportionately, but more likely indicated their sequential use. Nest temperatures may play a key role in energy use, with cooler temperature nests increasing the overall energy demands placed on developing embryos.

In 2006, we tracked a single Green Turtle after nesting Latakia, providing the first evidence of foraging grounds and migratory routes from this rookery which is one of the most important nesting areas in the Mediterranean. Tracking lasted 95 days during which time the turtle migrated southward (minimum average speed=0.9 km/h) and then westward (minimum average speed=1.6 km/h), following the coast. The turtle settled in the Bay of As Sallum, western Egypt where it remained in neritic, coastal waters for the final 26 days of transmissions. These results, when combined with findings from Cyprus, Turkey and Greece, further highlight the importance of the North African coast for foraging adult turtles and the need for effective conservation measures to be adopted there.

Adult sex ratio is a crucial parameter in population dynamics, with environmental sex determination being of particularly high theoretical and conservation interest. Monitoring hatchling sex ratio over time and space and understanding its variation may prove essential for effective con- servation of populations of species with temperature-dependent sex determination. We used clutch incubation duration to estimate sex ratio of hatchlings produced on Zakynthos (Greece), the largest rookery of loggerhead sea turtles Caretta caretta (a species with temperature-dependent sex deter- mination: colder temperatures producing more males) in the Mediterranean. We also explored spa- tial variation in hatchling sex ratio estimates within this nesting area. The estimated hatchling sex ratio on Zakynthos was female-biased in 2002 and 2003 (68 and 75% females estimated, respec- tively). We found significant differences in incubation durations between the 6 nesting beaches of Zakynthos, which result in pronounced differences in estimated hatchling sex ratio. Little is known about the relationship between adult and hatchling sex ratio or about the minimum number of males needed in healthy sea turtle populations. With global climate change, however, particular attention should be paid to the protection of the beaches that are apparently characterised by comparatively cold incubation temperatures to provide females with a variety of incubation environments.

Mating systems are a central component in the evolution of animal life histories and in conservation genetics. The patterns of male reproductive skew and of paternal shares in batches of offspring, for example, affect genetic effective population size. A prominent characteristic of mating systems of sea turtles seem to be a considerable intra- and interspecific variability in the degree of polyandry. Because of the difficulty of observing the mating behaviour of sea turtles directly in the open sea, genetic paternity analysis is particularly useful for gaining insights into this aspect of their reproductive behaviour. We investigated patterns of multiple paternity in clutches of loggerhead sea turtles in the largest Mediterranean rookery using four highly variable microsatellite loci. Furthermore, we tested for a relationship between the number of fathers detected in clutches and body size of females. More than one father was detected in the clutches of 14 out of 15 females, with two clutches revealing the contribution of at least five males. In more than half the cases, the contributions of different fathers to a clutch did not depart from equality. The number of detected fathers significantly increased with increasing female body size. This relationship indicates that males may prefer to mate with large, and therefore productive, females. Our results suggest that polyandry is likely to increase effective population size compared to a population in which females would mate with only one male; male reproductive contributions being equal.

Female sea turtles typically lay several clutches during each nesting season and rest in the time between laying clutches (the internesting period) in the waters off the nesting beach. Adequate protection of turtles in the internesting area requires knowledge on their spatial behaviour and was so far hampered by methodological limitations. Satellite telemetry data of exceptionally high quality allowed us to scrutinize internesting area use of four loggerhead sea turtles nesting in the Bay of Laganas (Zakynthos, Greece). We assessed the efficacy of two zones of a marine reserve with distinct levels of protection (boats are excluded from one zone). Most of the obtained locations of three of the four turtles were within the bay, a result consistent with a strategy to minimize energy expenditure. Turtles showed no obvious preference for the highly protected area of the bay. The availability of warmer water in the less protected area may be more crucial than avoidance of boat disturbance.

The sandy beaches of Zakynthos Island support the largest single nesting aggregation in the Mediterranean Region of the endangered loggerhead turtle Caretta caretta. The present study attempts to determine possible correlations between a series of habitat variables and nest site selection. Nesting activities, including total and nesting emergences were examined in response to the recorded biotic and abiotic variables. The results of the analysis indicate that beach width is the most critical habitat variable affecting nest site selection. Further analysis of nesting performance implies that sea turtles use multiple environmental cues for nest site selection during the different steps of the nesting processes such as emergence from the surf and nesting. Nevertheless, we caution that a detailed study needs to be conducted over a more extensive period of time to verify these suggestions.

In this paper, we evaluate the effectiveness of different well-tested conservation techniques in an effort to successfully protect sea turtle nests. From an eight-year study on the island of Zakynthos, West Greece, we have experimentally investigated the effectiveness of two different conservation techniques applied to loggerhead sea turtle nests and provided statistical measures to evaluate their conservation value. The categories of nests evaluated include: (i) nests incubated in situ, (ii) translocated into a beach hatchery, or (iii) protected by metal cages. Results of the analysis showed significant interannual variations in hatching success as obtained for each one of the three groups of nests. Significant differences were also observed when comparing hatching success data among the three groups of nests during the eight-year period. Overall, our results indicate that relocation of nests laid at highly threatened locations and the placement of protective cages on nests in situ provide adequate conservation measures that could allow an increase in hatchling production; although their choice and application should be based on the specific conditions and threats of each nest.

Offspring sex ratio is an important demographic parameter and, given its determination by incubation temperature in sea turtles, might be a key factor for their conservation under climate warming. An appealing approach to estimate hatchling sex ratios is to measure sand temperatures at nest depth and deduce hatchling sex ratios from a beforehand-established relationship of hatchling sex ratio and sand temperature. Such estimates will only be accurate though if metabolic heat produced by the embryos is considered. Judging whether metabolic heating has a potential effect on hatchling sex ratios without actually measuring temperature within clutches would greatly facilitate monitoring protocols. We tested for a relationship between the amount of metabolic heating and the number of developed embryos as well as clutch size in the largest known loggerhead sea turtle (Caretta caretta) population of the Mediterranean on Zakynthos (Greece). Temperatures were measured within 20 nests as well as at a reference site in the sand at nest depth. Metabolic heating was detected, but only during the last third of the incubation period did nests heat up considerably (1.6 °C on average) above the temperature of the surrounding sand. During the middle third of incubation, when sex is determined, the amount of metabolic heating was negligible. The amount of metabolic heating during the last third of the incubation duration was significantly correlated to the number of offspring developed to at least about 75% of incubation duration. This factor explained nearly 50% of variation in metabolic heating. Metabolic heating was also significantly correlated to clutch size. Given that clutch size within the Mediterranean is largest in Zakynthos loggerheads, we conclude that metabolic heating can be ignored in the estimate of hatchling sex ratios in Mediterranean loggerhead populations. These results thus provide the basis for a feasible monitoring of hatchling sex ratios in the loggerhead sea turtle in the Mediterranean.

Data on tag recoveries of 31 adult female and two juvenile loggerhead turtles (Caretta caretta) along the eastern Adriatic coast were analysed. All adult females originated from Mediterranean nesting areas: 29 from Greece, one from Cyprus and one from Turkey. This is the largest aggregation of long-distance tag recoveries in the Mediterranean for the loggerhead population nesting in Greece. The short period elapsed between the last observation of some turtles on the nesting beach and their recovery suggests the existence of a migratory pathway from the Ionian Sea into the Adriatic. Recoveries of some turtles within a period of a year indicate that loggerheads reside in the Adriatic for some time, suggesting that some individuals may demonstrate a level of site fidelity for these marine habitats. The high concentration of tag returns in the shallow northern Adriatic highlights this region as an important feeding habitat, for the population nesting in Greece in particular. The Ionian–Adriatic loggerheads form an important management sub-unit for this nesting population.

Pivotal temperature (the constant temperature giving 50% of each sex) for two clutches of loggerhead sea turtles (Caretta caretta) from Kyparissia Bay, Greece, was 29.3°C. Pivotal incubation duration (the time from laying to hatching giving 50% of each sex) was 52.6 days. These values are close to those obtained for this species in Brazil and the United States, providing further evidence that these characteristics are relatively conservative in different populations. Methodological differences between different experiments and limitations on accuracy of equipment make the detection of small differences problematic. Comparison of incubation durations in the field with the pivotal durations obtained here suggest that hatchling sex ratio on some Mediterranean beaches is female biased but probably varies considerably within this region.

Loggerhead turtles Caretta caretta use a wide variety of foraging strategies, and some populations forage in sub-optimal habitats. Different foraging strategies may not be equiva- lent in terms of fitness and may result in differences in adult body size and clutch size among populations. Accordingly, we tested whether differences in clutch size among rookeries in the Mediterranean Sea are related to differential use of foraging grounds of contrasting productivity. Stable isotope ratios of carbon and nitrogen of turtle hatchlings from 8 Mediterranean rookeries were used to characterise the foraging grounds of their mothers. Clutch size was also studied in each rookery to assess reproductive output linked to foraging ground productivity. According to stable isotope ratios, most of the females nesting in the considered rookeries foraged in the southern Ion- ian Sea. The highly productive Adriatic/northern Ionian Sea region was mainly used by females nesting in western Greece. The explanation for these patterns might be linked to water circulation patterns and drifting trajectories followed during developmental migrations, which might determine individual knowledge on the location of productive foraging patches. Average clutch size in each rookery was positively correlated to the proportion of females accessing highly productive areas such as the Adriatic/northern Ionian Sea. This has a strong influence on reproductive out-put, and hence females using the most productive foraging grounds had the largest clutch sizes. Fil: Cardona, Luis. Universidad de Barcelona; España Fil: Clusa, Marcel. Universidad de Barcelona; España Fil: Eder, Elena Beatriz. Universidad de Barcelona; España. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico; Argentina Fil: Demetropoulos, Andreas. Cyprus Wildlife Society; Chipre Fil: Margaritoulis, Dimitris. The Sea Turtle Protection Society of Greece; Grecia Fil: Rees, Alan F.. The Sea Turtle Protection Society of Greece; Grecia Fil: Hamza, Abdulmaula A.. Alfateh University; Libia Fil: Khalil, Mona. MEDASSET; Líbano Fil: Levy, Yaniv. Nature Parks Authority; Israel Fil: Türkozan, Oguz. Adnan Menderes University; Turquía Fil: Marín, Isabel. Universidad de Barcelona; España Fil: Aguilar, Alex. Universidad de Barcelona; España

Standardized ground surveys over six nesting seasons (1984–1989) along 42 km of beach at Kyparissia Bay, western Peloponnesus (Greece) revealed uneven spatial distribution in the nesting activity of the Loggerhead Turtle, Caretta caretta. Further surveys (1990–2000) at the most turtle-frequented part of the bay and the inclusion of a new beach sector have led to the determination of a 9.5 km core area where about 84% of the total nesting effort concentrates with an average nesting density of 60.6 nests/km/season. The average nesting effort along the bay is estimated at about 620 nests/season and therefore Kyparissia Bay is considered the second most turtle-frequented nesting area in the Mediterranean after Zakynthos. In the course of seventeen seasons (1984–2000) no distinguishable trend in the annual number of nests has been detected, albeit the great annual fluctuations (reaching 324%) may hide it. Although the duration of the nesting season (82.7 days on the average) seems similar to respectiv...

As the loggerhead turtle (Caretta caretta) is a philopatric species with a strong genetic structure, the analysis of mtDNA can be used to track evolutionary and colonisation events. In this study we use a genetic approach to understand the population structure of C. caretta in the Mediterranean Sea and to test whether loggerheads could have colonised the Mediterranean during the Pleistocene and survived the cold phases in warm refugia. We amplified a long mtDNA D-loop fragment (815. bp) from 168 dead hatchlings sampled from a selection of rookeries in the Eastern Mediterranean: Libya, Israel, Lebanon, Cyprus and Greece. Previously published data from Turkey and Calabria (Southern Italy) were also included in the analyses. The population nesting in Libya emerged as the oldest population in the Mediterranean, dating from the Pleistocene ca. 65,000. years ago (20,000-200,000). This reveals that the Libyan population might have settled in the Mediterranean basin before the end of the last glacial period. The remaining nesting sites, except Calabria, were subsequently colonised as the population expanded. The populations nesting in Eastern Turkey and Western Greece settled ca. 30,000. years ago (10,000-100,000), whereas the remaining populations originated as a result of a more recent Holocenic expansion. As Calabria presented a unique Atlantic haplotype, found nowhere else in the Mediterranean, we consider this nesting site as the result of an independent colonisation event from the Atlantic and not the recent spread of Mediterranean populations. This reveals that the current genetic structure of C. caretta rookeries in the Mediterranean would be the result of at least two colonisation events from the Atlantic, the oldest one in Libya and a most recent in Calabria, combined with local extinctions during Pleistocenic glaciations and re-colonisations from glacial refugia in Libya, Eastern Turkey and Western Greece. © 2012 Elsevier B.V.

To assess population genetic structure and evolutionary relationships among nesting populations of loggerhead turtles (Caretta caretta), we analyzed mitochondrial (mt) DNA variation in 113 samples from four nesting beaches in the northwestern Atlantic Ocean and from one nesting beach in the Mediterranean Sea. Significant differences in haplotype frequency between nesting populations in Florida and in Georgia/South Carolina, and between both of these assemblages and the Mediterranean nesting colony, indicate substantial restrictions on contemporary gene flow between regional populations, and therefore a strong tendency for natal homing by females. Nonetheless, this regional genetic structure appears shallow, indicating recent evolutionary connections among rookeries. Data from tag returns and mtDNA, as well as geological considerations, suggest that over short evolutionary time scales (perhaps a few thousand years), dispersal by female loggerheads is sufficient to allow colonization of appropriate habitat in proximity to established rookeries but is too low to significantly affect the population dynamics of rookeries on a contemporary time scale. These data indicate that nesting populations of the loggerhead turtle must be managed as demographically independent units. The population subdivisions based on mtDNA analyses are concordant with previously reported distinctions between Florida and Georgia/South Carolina nesting populations based on environmental markers, tag recaptures, and morphology. Resumen: Con el proposito de evaluar la estructura genetica de las poblaciones y las relaciones evolutivas entre poblaciones anidadoras de la tortuga caguama (Caretta caretta), analizamos la variacion en el ADN mitocondrial (ADNmt) de 113 muestras provenientes de cuatro playas de anidacion en el Oceano Atlantico noroccidental y de una en el Mar Mediterraneo. Las diferencias significativas en la frecuencia de haplotipos entre poblaciones anidadoras en Florida y en Georgia/Carolina del Sur, y entre estos dos grupos y la colonia anidadora en el Mediterraneo, indican limitaciones substanciales en el flujo genetico contemporaneo entre poblaciones regionales y, por lo tanto. una tendencia de las hembras a anidar en sus playas de nacimiento (“natal homing”). A pesar de lo anterior, dado que las distancias geneticas son pequenas se deducen conecciones evolutivas recientes entre colonias anidadoras. Los resultados de marcaje y del ADNmt, ademas de consideraciones geologicas, sugieren que a lo largo de escalas evolutivas pequenas (quiza unos cuantos miles de anos), la dispersion de caguamas hembras es suficiente para permitir la colonizacion de habitats apropiados en la proximidad de playas de anida cion establecidas pero es demasiado leve para afectar significativamente la dinamica de poblaciones de las colonias anidadoras en una escala de tiempo contemporanea. Nuestra informacion indica que las poblaciones anidadoras de la tortuga coguama deben ser manejadas como unidades demograficas independientes. La subdivision poblacional basada en los analisis del ADNmt concuerdan con las distinciones ya reportadas entre las poblaciones anidadoras en Florida y Georgia/Carolina del Sur basadas en marcadores ambientales. recapturas de tortugas marcadas y morfologa.

While telemetry is an invaluable tool for tracking animal movement patterns, the data generated by this technique is often challenging to interpret. Here, we addressed this issue by developing a novel method, based on changepoint analysis, which incorporated both the horizontal and vertical movement metrics and compared this output to that from a switching state-space model (SSSM) that categorized behavior based on horizontal movement metrics. We deployed 20 satellite transmitters on postnesting loggerhead turtles at Rethymno, Crete, Greece between 2010 and 2011 to monitor their at-sea behavior. We used both models to identify behavioral changes, such as the switches from migration to foraging, and from foraging to overwintering. The satellite-tracked turtles exhibited three discrete migratory strategies, with 9 turtles migrating southwards to the coast of northern Africa, 6 turtles migrating northwards into the Aegean Sea, and 4 turtles remaining resident in the waters of Crete. The SSSM readily identified the switch from transiting to ARS behavior in most animals, but the CPA model was able to distinguish multiple modes and more subtle shifts in behavior corresponding with shifts from migration to foraging to overwintering behaviors. We have shown that by incorporating vertical movement metrics into the analysis of telemetry data, previously hidden shifts in behavior can be revealed. The resulting increase in ability to discern complex behavioral patterns of animals remotely will likely yield better management and conservation decisions for a wide array of organisms.