Determinate or indeterminate growth? Revisiting the growth strategy of sea turtles

This is the author's accepted manuscript.The final version is available from Inter Research via the DOI in this record.Traditionally, growth can be either determinate, ceasing during the natural lifespan of individuals, or indeterminate, persisting throughout life. Although indeterminate growth is a widely accepted strategy and believed to be ubiquitous among long-lived species, it may not be as common as previously thought. Sea turtles are believed to be indeterminate growers despite the paucity of long-term studies into post-maturity growth. In this study, we provide the first temporal analysis of post-maturity growth rates in wild living sea turtles, using 26 yr of data on individual measurements of females nesting in Cyprus. We used generalised additive/linear mixed models to incorporate multiple growth measurements for each female and model post-maturity growth over time. We found post-maturity growth to persist in green Chelonia mydas and loggerhead Caretta caretta turtles, with growth decreasing for approximately 14 yr before plateauing around zero for a further decade solely in green turtles. We also found growth to be independent of size at sexual maturity in both species. Additionally, although annual growth and compound annual growth rates were higher in green turtles than in loggerhead turtles, this difference was not statistically significant. While indeterminate growth is believed to be a key life-history trait of ectothermic vertebrates, here, we provide evidence of determinate growth in green and loggerhead turtles and suggest that determinate growth is a life-history trait shared by cheloniid species. Our results highlight the need for long-term studies to refine life-history models and further our understanding of ageing and longevity of wild sea turtles for conservation and management.Fieldwork was supported by the British Associate of Tortoise Keepers, British Chelonia Group, British High Commission in Cyprus, British Residents Society, Carnegie Trust for the Universities of Scotland, Darwin Initiative, Erwin Warth Foundation, Friends of SPOT, Glasgow Uni versity Court, Kuzey Kıbrıs Turkcell, MEDASSET UK, and Natural Environment Research Council

Shelf life: Neritic habitat use of a turtle population highly threatened by fisheries

This is the final version. Available on open access from Wiley via the DOI in this recordAim: It is difficult to mitigate threats to marine vertebrates until their habitat use is understood. We report on a decade of satellite tracking loggerhead turtles (Caretta caretta) from an important nesting site to determine priority habitats for their protection in a region where they are known to be heavily impacted by fisheries. Location: Cyprus, Eastern Mediterranean. Method: We tracked 27 adult female loggerheads between 2001 and 2012 from North Cyprus nesting beaches. To eliminate potential biases, we included females nesting on all coasts of our study area, at different periods of the nesting season and from a range of size classes. Results: Foraging sites were distributed over the continental shelf of Cyprus, the Levant and North Africa, up to a maximum distance of 2100 km from nesting sites. Foraging sites were clustered in (1) near-shore waters of Cyprus and Syria, (2) offshore waters of Egypt and (3) offshore and near-shore regions of Libya and Tunisia. The North Cyprus and west Egypt/east Libyan coasts are important areas for loggerhead turtles during migration. Movement patterns within foraging sites strongly suggest benthic feeding in discrete areas. Early nesters visited other rookeries in Turkey, Syria and Israel where they likely laid further clutches. Tracking suggests minimum annual mortality of 11%, comparable to other fishery-impacted loggerhead populations. Main conclusions: This work further highlights the importance of neritic habitats of Libya and Tunisia as areas likely used by loggerhead turtles from many of the Mediterranean rookeries and where the threat of fisheries bycatch is high. Our tracking data also suggest that anthropogenic mortalities may have occurred in North Cyprus, Syria and Egypt; all within near-shore marine areas where small-scale fisheries operate. Protection of this species across many geopolitical units is a major challenge and documenting their distribution is an important first step.Peoples Trust for Endangered SpeciesBritish Chelonia GroupUnited States Agency for International DevelopmentBP EgyptApacheNatural Environment Research Council (NERC)Erwin Warth FoundationKuzey Kıbrıs TurkcellEktam KıbrısSEATURTLE.orgMEDASSETDarwin InitiativeBritish High Commission in CyprusBritish Residents Society of North CyprusMarine Turtle Conservation ProjectMarine Turtle Research GroupSociety for the Protection of Turtles in North Cyprus (SPOT)North Cyprus Department of Environmental Protectio

Defining conservation units with enhanced molecular tools to reveal fine scale structuring among Mediterranean green turtle rookeries

This is the final version. Available on open access from Elsevier via the DOI in this recordData accessibility: the data associated with this article is avaiable in the Dryad Digital Repository: https://datadryad.org/stash/dataset/doi:10.5061/dryad.7db01Understanding the connectivity among populations is a key research priority for species of conservation concern. Genetic tools are widely used for this purpose, but the results can be limited by the resolution of the genetic markers in relation to the species and geographic scale. Here, we investigated natal philopatry in green turtles (Chelonia mydas) from four rookeries within close geographic proximity (~200 km) on the Mediterranean island of Cyprus. We genotyped hypervariable mtSTRs, a mtDNA control region sequence (CR) and 13 microsatellite loci to genetically characterise 479 green turtles using markers with different modes of inheritance. We demonstrated matrilineal stock structure for the first time among Mediterranean green turtle rookeries. This result contradicts previous regional assessments and supports a growing body of evidence that green turtles exhibit a more precise level of natal site fidelity than has commonly been recognised. The microsatellites detected weak male philopatry with significant stock structure among three of the six pairwise comparisons. The absence of Atlantic CR haplotypes and mtSTRs amongst these robust sample sizes reaffirms the reproductive isolation of Mediterranean green turtles and supports their status as a subpopulation. A power analysis effectively demonstrated that the mtDNA genetic markers previously employed to evaluate regional stock identity were confounded by an insufficient resolution considering the recent colonisation of this region. These findings improve the regional understanding of stock connectivity and illustrate the importance of using suitable genetic markers to define appropriate units for management and conservation.Natural Environment Research Council (NERC)Ministerio de Economía y Competitividad, Spai

The importance of passive integrated transponder (PIT) tags for measuring life-history traits of sea turtles

This is the final version. Available from the publisher via the DOI in this record.Capture-mark-recapture studies rely on the identification of individuals through time, using markers or tags, which are assumed to be retained. This assumption, however, may be violated, having implications for population models. In sea turtles, individual identification is typically based on external flipper tags, which can be combined with internal passive integrated transponder (PIT) tags. Despite the extensive use of flipper tags, few studies have modelled tag loss using continuous functions. Using a 26-year dataset for sympatrically nesting green (Chelonia mydas) and loggerhead (Caretta caretta) turtles, this study aims to assess how PIT tag use increases the accuracy of estimates of life-history traits. The addition of PIT tags improved female identification: between 2000 and 2017, 53% of green turtles and 29% of loggerhead turtles were identified from PIT tags alone. We found flipper and PIT tag losses were best described by decreasing logistic curves with lower asymptotes. Excluding PIT tags from our dataset led to underestimation of flipper tag loss, reproductive periodicity, reproductive longevity and annual survival, and overestimation of female abundance and recruitment for both species. This shows the importance of PIT tags in improving the accuracy of estimates of life-history traits. Thus, estimates where tag loss has not been corrected for should be interpreted with caution and could bias IUCN Red List assessments. As such, long-term population monitoring programmes should aim to estimate tag loss and assess the impact of loss on life-history estimates, to provide robust estimates without which population models and stock assessments cannot be derived accuratel

Detecting green shoots of recovery: The importance of long-term individual-based monitoring of marine turtles

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this recordPopulation monitoring is an essential part of evaluating the effectiveness of management interventions for conservation. Coastal breeding aggregations of marine vertebrate species that come ashore to pup or nest provide an opportunistic window of observation into otherwise widely dispersed populations. Green turtle (Chelonia mydas) nesting on the north and west coasts of northern Cyprus has been monitored consistently and exhaustively since 1993, with an intensive saturation tagging programme running at one key site for the same duration. This historically depleted nesting population is showing signs of recovery, possibly in response to nest protection approaching two decades, with increasing nest numbers and rising levels of recruitment. Strong correlation between year-to-year magnitude of nesting and the proportion of new breeders in the nesting cohort implies that recruitment of new individuals to the breeding population is an important driver of this recovery trend. Recent changes in fishing activities may be impacting the local juvenile neritic stage, however, which may hinder this potential recovery. Individuals returning to breed after two years laid fewer clutches than those returning after three or four years, demonstrating a trade-off between remigration interval and breeding output. Average clutch frequencies have remained stable around a median of three clutches a year per female despite the demographic shift towards new nesters, which typically lay fewer clutches in their first season. We show that where local fecundity has been adequately assessed, the use of average clutch frequencies can be a reliable method for deriving nester abundance from nest counts. Index sites where individual-based monitoring is possible will be important in monitoring long-term climate driven changes in reproductive rates.European Social Fun

Spatial ecology of loggerhead turtles: Insights from stable isotope markers and satellite telemetry

This is the final version. Available on open access from Wiley via the DOI in this recordAim 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.Natural Environment Research Council (NERC

Behavioural patterns, spatial utilisation and landings composition of a small-scale fishery in the eastern Mediterranean

This is the final version. Available from Elsevier via the DOI in this record. Data Availability: Data will be made available on request.Small-scale fisheries (SSFs) are crucial for global food security and cultural heritage, however, information on their spatial distribution and practices are often lacking, precluding effective management and mitigation of ecological impacts. This is acutely the case in the eastern Mediterranean basin, where, despite concerns being raised regarding the magnitude of marine turtle bycatch in SSFs over two decades ago, a poor understanding of the fishery persists. To address this knowledge gap, we characterised the SSF fleet of Northern Cyprus through a combination of onboard observations, fisher self-reporting and vessel tracking to provide the first comprehensive overview of the fishery. Northern Cyprus had a fleet size, standardised by coastline length, ranked 14th of 23 Mediterranean fleets assessed, with an estimated 49542 and 57198 fishing days in total in 2020 and 2021, respectively. Vessels operated mainly over the continental shelf (< 200 m) and were predominantly active during the night (53.2%, n = 573378 locations). Clear crepuscular peaks in vessel activity and gear deployment raise concerns over spatiotemporal overlap with vulnerable species, even within MPAs previously established to protect them. Fishers (n = 1296 fishing operations) predominantly utilised static and demersal gear types including gill nets (35.0%), trammel nets (27.3%), trammel and gill nets combined (20.3%), demersal longlines (17.0%) and handlines (0.5%). Landings composition was highly diverse with a minimum of 238 different taxa identified, including, but not limited to, 123 species of bony fish, 22 elasmobranch species, 3 marine turtle species and 12 mollusc species of which 18.6% are considered threatened either at a Mediterranean or global scale. However, over 70.0% of total landing mass was comprised of only five species including bogue (Boops boops), picarel (Spicara smaris), blotched picarel (Spicara maena), greater amberjack (Seriola dumerili) and Mediterranean parrotfish (Sparisoma cretense). As the most up to date and detailed understanding of this fishery's operating behaviours, our research compares the results obtained from onboard observer and self-reporting fisher sampling methodologies and discusses the caveats of each and identifies potential opportunities to adapt existing practices and MPAs to improve long-term sustainability of the fishery, whilst maintaining its socio-economic benefits to the local community.Mava FoundationUniversity of ExeterUniversity of ExeterCERECON projec

Diet-related selectivity of macroplastic ingestion in green turtles (Chelonia mydas) in the eastern Mediterranean

Understanding the drivers of key interactions between marine vertebrates and plastic pollution is now considered a research priority. Sea turtles are primarily visual predators, with the ability to discriminate according to colour and shape; therefore these factors play a role in feeding choices. Classifcation methodologies of ingested plastic currently do not record these variables, however here, refned protocols allow us to test the hypothesis that plastic is selectively ingested when it resembles the food items of green turtles (Chelonia mydas). Turtles in the eastern Mediterranean displayed strong diet-related selectivity towards certain types (sheet and threadlike), colours (black, clear and green) and shapes (linear items strongly preferred) of plastic when compared to the environmental baseline of plastic beach debris. There was a signifcant negative relationship between size of turtle (curved carapace length) and number/mass of plastic pieces ingested, which may be explained through naivety and/or ontogenetic shifts in diet. Further investigation in other species and sites are needed to more fully ascertain the role of selectivity in plastic ingestion in this marine vertebrate group

Dietary analysis of two sympatric marine turtle species in the eastern Mediterranean

This is the final version. Available from Springer via the DOI in this record. Data and any available visuals, such as figures and tables, will be provided upon reasonable request to the corresponding author. Any codes created to assist in data analysis and/or any software application utilised in the current study will be provided upon reasonable request submitted to the corresponding author.Dietary studies provide key insights into threats and changes within ecosystems and subsequent impacts on focal species. Diet is particularly challenging to study within marine environments and therefore is often poorly understood. Here, we examined the diet of stranded and bycaught loggerhead (Caretta caretta) and green turtles (Chelonia mydas) in North Cyprus (35.33° N, 33.47° E) between 2011 and 2019. A total of 129 taxa were recorded in the diet of loggerhead turtles (n = 45), which were predominantly carnivorous (on average 72.1% of dietary biomass), foraging on a large variety of invertebrates, macroalgae, seagrasses and bony fish in low frequencies. Despite this opportunistic foraging strategy, one species was particularly dominant, the sponge Chondrosia reniformis (21.5%). Consumption of this sponge decreased with increasing turtle size. A greater degree of herbivory was found in green turtles (n = 40) which predominantly consumed seagrasses and macroalgae (88.8%) with a total of 101 taxa recorded. The most dominant species was a Lessepsian invasive seagrass, Halophila stipulacea (31.1%). This is the highest percentage recorded for this species in green turtle diet in the Mediterranean thus far. With increasing turtle size, the percentage of seagrass consumed increased with a concomitant decrease in macroalgae. Seagrass was consumed year-round. Omnivory occurred in all green turtle size classes but reduced in larger turtles (> 75 cm CCL) suggesting a slow ontogenetic dietary shift. Macroplastic ingestion was more common in green (31.6% of individuals) than loggerhead turtles (5.7%). This study provides the most complete dietary list for marine turtles in the eastern Mediterranean.European CommissionErwin Warth FoundationKarşıyaka Turtle WatchAngela WadsworthMaureen and Tony HutchinsonKuzey Kıbrıs TurkcellMAVA FoundationNatural Environment Research Council (NERC)University of Exete

Network analysis of sea turtle movements and connectivity: A tool for conservation prioritization

This is the final version. Available on open access from Wiley via the DOI in this recordData availability statement: The data that support the findings of this study are available in the Supplementary Material of this article and Zenodo (https://doi.org/10.5281/zenodo.5898578). Details for all animals included in this study are provided in Appendices S1 and S2. Data used to create the spatial networks are listed in the Appendices S3 and S4. The geospatial files for all networks are available on the Migratory Connectivity in the Ocean Project website (https://mico.eco) and Dryad (https://doi.org/10.5061/dryad.j3tx95xg9). Additional data that support the findings of this study are available from the corresponding author upon reasonable request.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.International Climate Initiative (IKI)German Federal Ministry for the Environment, Nature Conservation and Nuclear Safety (BMU