Feeding ecology of the green turtle (Chelonia mydas) at rocky reefs in western South Atlantic

Feeding ecology of green turtles was investigated between January 2005 and April 2008 at Arvoredo Reserve, Brazil (27°17′S, 48°18′W). Data were obtained through the performance of observational sessions, geo-referenced counts, benthic surveys, capture and recapture of individuals, and oesophageal lavages. This protected area was identified as an important green turtle feeding ground, used year-round by juveniles (curved carapace length = 32–83 cm). Turtles fed close to the rocky shores of the area and selected grazing sites commonly at hard-to-reach, near-vertical portions of the rocks. They were less active in cold months, and more abundant at shallow areas of the reef (0–5 m), where their preferred food items occurred. Their diet was dominated by macroalgae species but invertebrates were also present. Their main food item was the red algae Pterocladiella capillacea, which seems to be eaten through periodical cropping of its tips. Observational methods such as the ones applied here could be incorporated to other research programs aiming to understand the relationships between Chelonia mydas feeding populations and their environment.     

Flexible foraging movements of leatherback turtles across the North Atlantic Ocean

Some marine species have been shown to target foraging at particular hotspots of high prey abundance. However, we show here that in the year after a nesting season, female leatherback turtles (Dermochelys coriacea) in the Atlantic generally spend relatively little time in fixed hotspots, especially those with a surface signature revealed in satellite imagery, but rather tend to have a pattern of near continuous traveling. Associated with this traveling, distinct changes in dive behavior indicate that turtles constantly fine tune their foraging behavior and diel activity patterns in association with local conditions. Switches between nocturnal vs. diurnal activity are rare in the animal kingdom but may be essential for survival on a diet of gelatinous zooplankton where patches of high prey availability are rare. These results indicate that in their first year after nesting, leatherback turtles do not fit the general model of migration where responses to resources are suppressed during transit. However, their behavior may be different in their sabbatical years away from nesting beaches. Our results highlight the importance of whole-ocean fishing gear regulations to minimize turtle bycatch.     

Delayed ontogenic dietary shift and high levels of omnivory in green turtles (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) from the NW coast of Africa

Young green turtles (Chelonia mydas) spend their early lives as oceanic omnivores with a prevalence of animal prey. Once they settle into neritic habitats (recruitment), they are thought to shift rapidly to an herbivorous diet, as revealed by studies in the Greater Caribbean. However, the precise timing of the ontogenic dietary shift and the actual relevance of animal prey in the diet of neritic green turtles are poorly known elsewhere. Stable isotopes of carbon, sulfur and nitrogen in the carapace scutes of 19 green turtles from Mauritania (NW Africa), ranging from 26 to 102 cm in curved carapace length (CCLmin), were analyzed to test the hypothesis of a rapid dietary shift after recruitment. Although the length of residence time in neritic habitats increased with turtle length, as revealed by a significant correlation between turtle length and the δ¹³C and the δ³⁴S of the scutes, comparison of the δ¹⁵N of the innermost and outermost layers of carapace scutes demonstrated that consumption of macrophytes did not always start immediately after recruitment, and turtles often resumed an animal-based diet after starting to graze on seagrasses. As a consequence, seagrass consumption did not increase gradually with turtle size and animal prey largely contributed to the diet of turtles within the range 29–59 cm CCLmin (76–99% of assimilated nutrients). Seagrass consumption by turtles larger than 59 cm CCLmin was higher, but they still relied largely on animal prey (53–76% of assimilated nutrients). Thus, throughout most of their neritic juvenile life, green turtles from NW Africa would be better classified as omnivores rather than herbivores. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Can leftovers from predators be reliably used to monitor marine turtle hatchling sex-ratios? The implications of prey selection by ghost crabs

In marine turtles, the sex of an individual is determined by temperatures experienced during embryonic development. Gonad histological observation is still the only reliable way to determine hatchling sex, hampering the study of reproduction and of the demographic consequences of context-dependent sex-ratios, a subject of interest in a warming planet. We investigated whether hatchling remains from predation by Ocypode cursor can be used to estimate sex-ratio trends in a green turtle rookery at Poilão, Guinea-Bissau (10°52′N, 15°43′W). Sex could be readily determined in 77 and 79% of the predated hatchlings in 2008 and 2009, respectively. By comparing hatchlings killed by crabs, hatchlings accidentally dying on the reefs, and live hatchlings, we show that ghost crabs select the smaller prey, but do not select according to hatchling sex, which is explained by the lack of hatchling size dimorphism in this population. The proportion of male hatchlings was 0.45 ± 0.06 and 0.15 ± 0.06 for early and late-season clutches, respectively, these differences most likely being explained by rainfall. Using leftovers from predation by crabs may be a good solution to non-invasively monitor broad trends in sex-ratios of sea turtles.     

Foraging habitats and migration corridors utilized by a recovering subpopulation of adult female loggerhead sea turtles: implications for conservation

From 1998 to 2008, 68 adult female loggerhead sea turtles (Caretta caretta) were instrumented with platform transmitter terminals at nesting beaches in Georgia, North Carolina (NC) and South Carolina (SC) on the East Coast of the United States of America (30°48′N, 81°28′W to 33°51′N, 77°59′W). The majority of post-nesting loggerheads (N = 42, 62 %) migrated to foraging habitats in the Mid-Atlantic Bight during May–October, with a subsequent migration occurring during November–March to foraging habitats south of Cape Hatteras, NC. Nine (13 %) loggerheads initially foraged in the near-shore, coastal areas of the South Atlantic Bight, but moved to offshore habitats—closer to the Gulf Stream—during November–March, while fourteen (21 %) loggerheads remained in foraging areas along the mid-continental shelf off of the eastern coast of Florida and/or continued southward to Florida Bay and the Bahamas. The present study delineates important, post-nesting foraging habitats and migration corridors where loggerheads may interact with commercial fisheries—providing managers opportunities to develop and implement optimally effective conservation actions for the recovery of this threatened species.     

Feeding rates and prey selectivity of planktonic decapod larvae in the Western English Channel

Meroplankton are seasonally important contributors to the zooplankton, particularly at inshore sites, yet their feeding ecology is poorly known relative to holoplankton. While several studies have measured feeding in decapod larvae, few studies have examined the feeding rates of decapod larvae on natural prey assemblages throughout the reproductive season. We conducted 8 feeding experiments with Necora puber, Liocarcinus spp. and Upogebia spp. zoea larvae collected from the L4 monitoring site off Plymouth (50°15.00′N, 4°13.02′W) during spring–summer 2009 and 2010. This period spanned moderate-to-high food availability (0.5–1.6 µg chl-a L^?1), but a great range in food composition with small cells <20 µm dominating in 2010. Daily rations averaged 17, 60 and 22 % of body C for the 3 respective decapod species. Clearance rates differed according to prey type, and all 3 decapod genera showed evidence of selection of dinoflagellates. Importantly, small cells including nano- and pico-plankton were ingested, this being demonstrated independently by flow cytometric analysis of the feeding experiments and molecular analysis. PCR-based analysis of the haptophyte portion of the diet revealed ingestion of Isochrysis galbana by decapod larvae in the bottle incubations and Isochrysis galbana and Phaeocystis globosa by decapod larvae collected directly from the field. This study has shown that pico- and nano-sized plankton form an important supplement to the diverse and variable diet of decapod larvae.     

Home range of immature green turtles tracked at an offshore tropical reef using automated passive acoustic technology

Seventeen immature green turtles Chelonia mydas were tracked concurrently by automated ultrasonic receivers at a coral reef off North-Eastern Australia (September–December 2010, 16.4°S, 145.6°E). The majority (n = 11) were tracked for the entire 100-day study, the remainder for 23–85 days. Detection data aggregated at 30-min intervals produced median 6.5–35 daily locations for individual turtles. Home range areas (95 % utilisation distribution) were ≤1 km², $ {\bar{\text{x}}} $  ± SD = 0.74 km² ± 0.159. To the best of our knowledge, these are the first home range estimates for C. mydas foraging at offshore tropical reefs. The findings are important for conservation in revealing near-continuous presence of the same individuals within a small geographic area. Time between detections was very short (median <3 min) demonstrating passive ultrasonic technology can track multiple turtles in a foraging environment with higher temporal resolution than typically achieved by satellite tracking.     

Hitchhikers reveal cryptic host behavior: new insights from the association between Planes major and sea turtles in the Pacific Ocean

Studies that incorporate information from habitat-specific ecological interactions (e.g., epibiotic associations) can reveal valuable insights into the cryptic habitat-use patterns and behavior of marine vertebrates. Sea turtles, like other large, highly mobile marine vertebrates, are inherently difficult to study, and such information can inform the implementation of conservation measures. The presence of epipelagic epibionts, such as the flotsam crab Planes major, on sea turtles strongly suggests that neritic turtles have recently occupied epipelagic habitats (upper 200 m in areas with >200 m depth) and that epipelagic turtles spend time at or near the surface. We quantified the effects of turtle species, turtle size, and habitat (neritic or epipelagic) on the frequency of epibiosis (F ₀) by P. major on sea turtles in the Pacific Ocean. In neritic habitats, we found that loggerhead (F ₀ = 27.6 %) and olive ridley turtles (F ₀ = 26.2 %) host crabs frequently across a wide range of body sizes, and green turtles almost never host crabs (F ₀ = 0.7 %). These results suggest that loggerheads and olive ridleys display variable/flexible epipelagic-neritic transitions, while green turtles tend to transition unidirectionally at small body sizes. In epipelagic habitats, we found that loggerheads host crabs (F ₀ = 92.9 %) more frequently than olive ridleys (F ₀ = 50 %) and green turtles (F ₀ = 38.5 %). These results suggest that epipelagic loggerheads tend to spend more time at or near the surface than epipelagic olive ridleys and green turtles. Results of this study reveal new insights into habitat-use patterns and behavior of sea turtles and display how epibiont data can supplement data from more advanced technologies to gain a better understanding of the ecology of marine vertebrates during cryptic life stages.     

Temporal variability of deep-sea zooplankton in the Arabian Sea

Mesozooplankton samples from two stations in the Arabian Sea (WAST, 4,050 m, 16°15′N, 60°20′E; CAST, 3,950 m, 14°30′N, 64°30′E) were collected from the surface down to 20 m above bottom during three monsoon periods: the autumn intermonsoon in October 1995, the spring intermonsoon in April 1997, and the NE monsoon in February 1998. The main goal of this study is to enhance our knowledge on the effect of spatial and temporal differences in primary production and particle flux rates on the abundance and distribution of mesozooplankton, with special attention to the deep sea. Literature data indicate episodically high rates of primary production and particle flux in the region during the SW monsoon and the autumn intermonsoon. Set in this context, the zooplankton showed an in-phase coupling in biomass and abundance with the primary production in the surface 150 m. In the mesopelagic realm (150–1,050 m), the seasonal coupling was less clear. In the bathypelagic zone, below 1,050 m, there was no evidence of in-phase coupling, though temporal differences in the distribution of zooplankton abundance and biomass with depth between seasons could be shown by an analysis of covariance and an a posteriori test. The results suggest that the bathypelagic community responds to increased particle flux rates, but with longer time gaps than in the epipelagic zone. This is probably due to longer development and response times of zooplankton in the cold, deep-water environment, independent of possible lateral advection processes.     

Ecology of loggerhead marine turtles Caretta caretta in a neritic foraging habitat: movements, sex ratios and growth rates

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 (<2.7 cm yr^?1) that slowed with increasing body size. We revealed that an unexpectedly high proportion of animals were male (>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.     

Effects of ocean acidification, temperature and nutrient regimes on the appendicularian Oikopleura dioica: a mesocosm study

Increasing pCO₂ is hypothesized to induce shifts in plankton communities toward smaller cells, reduced carbon export rates and increased roles of gelatinous zooplankton. Appendicularians, among the most numerous pan-global “gelatinous” zooplankton, continuously produce filter-feeding houses, shortcutting marine food webs by ingesting submicron particles, and their discarded houses contribute significantly to carbon fluxes. We present a first mesocosm-scale study on the effects of temperature, pCO₂ and bloom structures on the appendicularian, Oikopleura dioica. There were effects of temperature and nutrients on phytoplankton communities. No shifts in functional phytoplankton groups, nor changes in particle sizes/morphotypes, known to impact appendicularian feeding, were observed under manipulated pCO₂ conditions. However, appendicularian abundance was positively correlated with increased pCO₂, temperature and nutrient levels, consistent with hypotheses concerning gelatinous zooplankton in future oceans. This suggests appendicularians will play more important roles in marine pelagic communities and vertical carbon transport under projected ocean acidification and elevated temperature scenarios.     

Foraging areas differentially affect reproductive output and interpretation of trends in abundance of loggerhead turtles

Diet items and habitat constitute some of the environmental resources that may be used differently by individuals within a population. Long-term fidelity by individuals to particular resources exemplifies individual specialization, a phenomenon that is becoming increasingly recognized across a wide range of species. Less is understood about the consequences of such specialization. Here, we investigate the effects of differential foraging ground use on reproductive output in 183 loggerhead sea turtles (Caretta caretta) nesting at Wassaw Island, Georgia (31.89°N, 80.97°W), between 2004 and 2011 with resulting possible fitness effects. Stable isotope analysis was used to assign the adult female loggerheads to one of three foraging areas in the Northwest Atlantic Ocean. Our data indicate that foraging area preference influences the size, fecundity, and breeding periodicity of adult female loggerhead turtles. We also found that the proportion of turtles originating from each foraging area varied significantly among the years examined. The change in the number of nesting females across the years of the study was not a result of uniform change from all foraging areas. We develop a novel approach to assess differential contributions of various foraging aggregations to changes in abundance of a sea turtle nesting aggregation using stable isotopes. Our approach can provide an improved understanding of the influences on the causes of increasing or decreasing population trends and allow more effective monitoring for these threatened species and other highly migratory species.     

Home range and habitat use of juvenile Atlantic green turtles (Chelonia mydas L.) on shallow reef habitats in Palm Beach, Florida, USA

Many animals, including sea turtles, alter their movements and home range in relation to the particular type and quality of the habitat occupied. When sufficient resources are available, individuals may develop affinities to specific areas for activities, such as foraging and (or) resting. In the case of green sea turtles (Chelonia mydas L.), after a number of years in the open ocean, juveniles recruit to shallow-water developmental habitats where they occupy distinct home ranges as they feed and grow to maturity. Our goal was to study the habitat use and home range movements of juvenile green turtles along a shallow, worm-rock reef tract in Palm Beach, Florida. Six turtles, measuring from 27.9 to 48.1 cm in straight carapace length and from 7.2 to 12.6 kg in mass, were tracked via ultrasonic telemetry from August to November 2003. Upon capture, each turtle’s esophagus was flushed via lavage to determine recently ingested foods. In addition, four turtles were recaptured and fitted with a time-depth recorder to study dive patterns. Home range areas measured with 100% minimum convex polygon and 95% fixed kernel estimators varied from 0.69 to 5.05 km² (mean=2.38±1.78 km²) and 0.73 to 4.89 km² (mean=2.09±1.80 km²), respectively. Home ranges and core areas of turtles were largely restricted to the reef tract itself, and showed considerable overlap between food and shelter sites. The mean number of dives during daylight hours (0600–1800 hours) was 84±5.0 dives, while the mean during night hours (1800–0600 hours) was 39±3.0 dives. Dives during the day were shallower (mean=3.20±1.26 m) than dives at night (mean=5.59±0.09 m). All six turtles were found to have a mixed diet of similar macroalgae and sponge fragments. Our results reveal that juvenile green turtles occupy stable home ranges along the nearshore worm-rock reefs of Southeast Florida, during the summer and fall. Determining which habitats are used by green turtles will assist conservation managers in their global effort to protect this endangered species.     

Satellite telemetry reveals behavioural plasticity in a green turtle population nesting in Sri Lanka

Satellite transmitters were deployed on ten green turtles (Chelonia mydas) nesting in Rekawa Sanctuary (RS-80.851°E 6.045°N), Sri Lanka, during 2006 and 2007 to determine inter-nesting and migratory behaviours and foraging habitats. Nine turtles subsequently nested at RS and demonstrated two inter-nesting strategies linked to the location of their residence sites. Three turtles used local shallow coastal sites within 60 km of RS during some or all of their inter-nesting periods and then returned to and settled at these sites on completion of their breeding seasons. In contrast, five individuals spent inter-nesting periods proximate to RS and then migrated to and settled at distant (>350 km) shallow coastal residence sites. Another turtle also spent inter-nesting periods proximate to RS and then migrated to a distant oceanic atoll and made forays into oceanic waters for 42 days before transmissions ceased. This behavioural plasticity informs conservation management beyond protection at the nesting beach.     

Determination of δ13C and δ15N and trophic fractionation in jellyfish: implications for food web ecology

Application of stable isotope analysis (SIA) in jellyfish allows definition of trophic patterns not detectable using gut content analysis alone, but analytical protocols require standardization to avoid bias in interpreting isotopic data. We determined δ¹³C and δ¹⁵N in Aurelia sp. from the northern Gulf of Mexico (30°00′N, 89°00′W–30°24′N, 88°00′W) to define differences in stable isotope composition between body parts and whole body, the effect of lipid extraction on δ¹³C in tissues, and fractionation values from medusa to prey. The isotopic composition of bell and whole Aurelia sp. was not different. The increase in δ¹³C values after lipid removal suggested a correction is needed. To aid future analyses, we derived a correction equation from empirical data for jellyfish samples. Laboratory feeding experiments indicated medusae increased +4 ‰ in δ¹³C and +0.1 ‰ in δ¹⁵N compared to their diet. These results suggest protocols commonly applied for other species may be inaccurate to define Aurelia sp. trophic ecology. Because Aurelia spp. are commonly found in marine ecosystems, accurately defining their trophic role by use of SIA has implications for understanding marine food webs worldwide.     

Growth and bleaching of the coral Oculina patagonica under different environmental conditions in the western Mediterranean Sea

The success that the putative alien species Oculina patagonica is able to survive under different environmental conditions may be benefiting its establishment and spreading along the Mediterranean Basin. Our objectives were to determine the response of this species, in terms of growth and bleaching, under different environmental conditions. Field data on colony growth and bleaching were obtained for a period of 18 months (from June 2010 to December 2011), in the Alicante Harbour (38°20′11″N, 00°29′11″W) and the Marine Protected Area of Tabarca (38°09′59″N, 00°28′56″W). Additionally, data on sedimentation rates, chlorophyll a concentration and organic matter were also collected. Moreover, the role of light over growth and bleaching of the coral was also studied with a field experiment. Our results showed that growth rates were similar among localities (eutrophic and oligotrophic environments), decreasing with increasing perimeter of the colony. Growth rates were at a minimum during cold months (13 °C) and enhanced during warm months until a threshold temperature; thereafter, bleaching was observed (>28 °C), being adverse for coral growth. In addition, light attenuation could act such as local stress, increasing the coral bleaching with the increase in seawater temperature. Our findings confirmed that O. patagonica has a broad tolerance to seawater temperature, irradiance and trophic water conditions, in addition to its ability to thrive through bleaching events, mainly in eutrophic environments, probably related to food availability.     

Stalked tunicates Boltenia ovifera form biogenic habitat in the rocky subtidal zone of Nova Scotia

The stalked tunicate Boltenia ovifera is widely distributed in the Arctic and North Atlantic oceans on rocky substrata at 10–300 m depth, although its ecological role in benthic communities is poorly understood. The distribution and abundance of B. ovifera were recorded at 10–100 m depth in towed-video transects in November 2011 and in June, July, and November 2012 off a wave-exposed headland near Halifax, Nova Scotia. Specimens also were collected at 30 m depth using SCUBA (44°26.88′N, 63°31.59′W) in September 2012. Areas dominated by B. ovifera had densities of 10–100 individuals m^?2 on rocky substrata at 30–60 m depth. These tunicate beds often were associated with sparse kelp (Agarum clathratum) and turf-forming red algae. A generalized additive model indicated that depth, substrate type, and benthic algal type were strong predictors of tunicate abundance. Twenty-two epibiotic species were found on specimens of B. ovifera, including juvenile conspecifics. Filter-feeding macroinvertebrates, including anemones and soft corals, were more abundant in areas with B. ovifera than in areas without these tunicates. Our findings suggest that beds of B. ovifera can act as biogenic habitat to enhance local species richness in the rocky subtidal zone of Nova Scotia.     

Modelling the effect of fibropapilloma disease on the somatic growth dynamics of Hawaiian green sea turtles

The effect of the tumour-forming disease, fibropapillomatosis, on the somatic growth dynamics of green turtles resident in the Pala’au foraging grounds (Moloka’i, Hawai’i) was evaluated using a Bayesian generalised additive mixed modelling approach. This regression model enabled us to account for fixed effects (fibropapilloma tumour severity), nonlinear covariate functional form (carapace size, sampling year) as well as random effects due to individual heterogeneity and correlation between repeated growth measurements on some turtles. Somatic growth rates were found to be nonlinear functions of carapace size and sampling year but were not a function of low-to-moderate tumour severity. On the other hand, growth rates were significantly lower for turtles with advanced fibropapillomatosis, which suggests a limited or threshold-specific disease effect. However, tumour severity was an increasing function of carapace size—larger turtles tended to have higher tumour severity scores, presumably due to longer exposure of larger (older) turtles to the factors that cause the disease. Hence turtles with advanced fibropapillomatosis tended to be the larger turtles, which confounds size and tumour severity in this study. But somatic growth rates for the Pala’au population have also declined since the mid-1980s (sampling year effect) while disease prevalence and severity increased from the mid-1980s before levelling off by the mid-1990s. It is unlikely that this decline was related to the increasing tumour severity because growth rates have also declined over the last 10–20 years for other green turtle populations resident in Hawaiian waters that have low or no disease prevalence. The declining somatic growth rate trends evident in the Hawaiian stock are more likely a density-dependent effect caused by a dramatic increase in abundance by this once-seriously-depleted stock since the mid-1980s. So despite increasing fibropapillomatosis risk over the last 20 years, only a limited effect on somatic growth dynamics was apparent and the Hawaiian green turtle stock continues to increase in abundance.     

Composition of green turtle feeding aggregations along the Japanese archipelago: implications for changes in composition with current flow

In order to develop effective conservation strategies for endangered migratory species, the link between feeding and breeding grounds needs to be clarified. In this study, the genetic compositions of consecutive Japanese feeding aggregations of green turtles (Chelonia mydas) along the Kuroshio Current were examined by mixed-stock analyses of mitochondrial DNA control-region sequences. The results indicated that the southern feeding aggregation around Yaeyama (24.3°N, 124.0°E) was sourced from various Pacific rookeries in the Yaeyama, Ogasawara, Western Pacific, and Indian Oceans and Southeast Asia. Among northern feeding aggregations, the Ginoza (26.5°N, 128.0°E) aggregation was also sourced from the Western Pacific Ocean, but the Nomaike (31.4°N, 130.1°E), Muroto (33.2°N, 134.2°E), and Kanto (35.6°N, 140.5°E) aggregations were contributed mostly by the closer Ogasawara rookeries. The reduced contribution from tropical Pacific rookeries to northern feeding aggregations and the significant correlation between genetic differentiation and geographical distance matrices of feeding aggregations indicated that most hatchlings from these regions transported by the Kuroshio Current settle in upstream feeding grounds along the Japanese archipelago, implying that current flow influences the composition of feeding aggregations. Differences in the composition of relatively close neritic feeding aggregations have important conservation implications, for which both regional and multinational conservation strategies are needed.