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Satellite telemetry and stable isotope analysis were used to confirm that oceanic areas (where water depths are >200 m) are alternative feeding habitats for adult female green sea turtles (Chelonia mydas), which have been thought to be obligate herbivores in neritic areas (where depths are <200 m). Four females were tagged with satellite transmitters and tracked during post-nesting periods from Ogasawara Islands, Japan. Three females migrated to neritic habitats, while transmissions from another female ceased in an oceanic habitat. The overall mean nighttime dive depths during oceanic swimming periods in two females were <20 m, implying that the main function of their nighttime dives were resting with neutral buoyancy, whereas the means in two other females were >20 m, implying that they not only rested, but also foraged on macroplankton that exhibit diel vertical migration. Comparisons of stable carbon and nitrogen isotope ratios between 89 females and the prey items in a three-source mixing model estimated that 69% of the females nesting on Ogasawara Islands mainly used neritic habitats and 31% mainly used oceanic habitats. Out of four females tracked by satellite, two females were inferred from isotope ratios to be neritic herbivores and the two others oceanic planktivores. Although post-nesting movements for four females were not completely consistent with the inferences from isotope ratios, possibly due to short tracking periods (28-42 days), their diving behaviors were consistent with the inferences. There were no relationships between body size and the two isotope ratios, indicating a lack of size-related differences in feeding habitat use by adult female green turtles, which was in contrast with loggerhead sea turtles (Caretta caretta). These results and previous findings suggest that ontogenetic habitat shifts by sea turtles are facultative, and consequently, their life histories are polymorphic.

Intrapopulation variation in habitat use is commonly seen among mobile animals, yet the mechanisms maintaining it have rarely been researched among untrackable species. To investigate how alternative life histories are maintained in a population of the loggerhead sea turtle (Caretta caretta), cumulative reproductive output was evaluated and compared between small planktivores inhabiting oceanic areas (with water depths > 200 m) and large benthivores inhabiting neritic areas (depths < 200 m) that sympatrically nested at Yakushima Island, Japan, from 1986 to 2011. In total, 362 nesting females sampled in three different years were classified into the two foraging groups based on stable isotope ratios in egg yolks. There were significant differences between the two foraging groups in most recorded life history parameters (clutch size, clutch frequency, breeding frequency, and remigration intervals), with the exception of emergence success. We did not find evidence of life history trade-offs, nor age-related changes in fecundity. Over the 26-year study period, we calculated a 2.4-fold greater reproductive output for neritic foragers than for oceanic ones, accounting for breeding and clutch frequency. Temporal consistencies in stable isotope ratios and remigration intervals within females suggested that female Japanese loggerheads show fidelity to respective foraging habitats throughout the adult stage. The large difference in productivity between the two groups was unlikely to be offset by the difference in survival during the period from aboveground emergence to first reproduction, suggesting that oceanic foragers have a lower level of fitness than neritic ones. Together with an absence of genetic structure between foraging groups, we infer that alternative life histories in a loggerhead turtle population are maintained by a conditional strategy.

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Information on population genetic structure is fundamental for recognizing management units of endangered species (Moritz 1994). Because population genetic structure depends on both the resolution and the inheritance modes of genetic markers, it should be analyzed through the combined use of multiple mark- ABSTRACT: Knowledge of detailed population genetic structure is crucial to conserve and manage endangered species effectively. Size-related variation in feeding-habitat use (neritic vs. oceanic) by adult loggerhead turtles Caretta caretta has been reported within several populations, and sympatric population subdivision was suspected. In the present study, genetic differences between the 2 feeding-habitat groups within 2 Japanese nesting sites were assessed, using 5 microsatellite loci and mitochondrial (mt) DNA sequences. There were no genotypic or haplotype differences between the feeding-habitat groups, which were defined by egg-yolk stable isotope ratios and body size, at both nesting sites, suggesting that both neritic and oceanic individuals belong to the same genetic population. Differences in feeding-habitat use are unlikely to be a limiting factor for gene flow between feeding-habitat groups and were thought to be the result of phenotypic plasticity rather than population subdivision. Gene flow among 5 nesting sites was assessed by pooling these feeding-habitat groups at each nesting site. Significant genetic structure by female natal homing was observed at the mtDNA level. However, no significant structure was found at the microsatellite DNA level, suggesting male-mediated gene flow caused by migration through courtship areas. Although nesting beaches are connected by male-mediated gene flow, which might have evolved as a mechanism to avoid genetic fragmentation by natal homing, extirpated beaches would not be easily recolonized from other nesting populations due to female philopatry. Therefore, conservation of individual nesting beaches is still needed to maintain the overall genetic diversity of Japanese loggerheads.KEY WORDS: Alternative life histories · Microsatellite DNA · Mitochondrial DNA · Phenotypic plasticity · Reptile · Caretta caretta Resale or republication not permitted without written consent of the publisherMar Ecol Prog Ser 424: [273][274][275][276][277][278][279][280][281][282][283] 2011 ers. Sequences of maternally inherited mitochondrial (mt) DNA and microsatellites of biparentally inherited nuclear DNA are the markers commonly used in recent wildlife population genetics (Frankham et al. 2002). In principle, due to their faster rate of evolution, the resolution of microsatellites is greater than that of mtDNA sequences. Nevertheless, in some cases, population differentiation inferred from mtDNA sequences is stronger than that from microsatellites (e.g. FitzSimmons et al. 1997b, Bowen et al. 2005, Carreras et al. 2007, Chen et al. 2008, Lukoschek et al. 2008, Okello et al. 2008, Caparroz et al. 2009, Hefti-Gautschi et al. 2009, Portnoy et al. 2010, and this has been attributed...

Intrapopulational polymorphism in habitat use is widely reported in many animal species. The phenomenon has recently also been recognized in adult female loggerhead sea turtles Caretta caretta, with small females tending to inhabit oceanic areas (where water depths are 4200 m) while presumably feeding pelagically and large females tending to inhabit neritic areas (where depths are o200 m) while presumably feeding benthically. In this study, dive recording satellite telemetry units were used to verify their foraging and diving behaviours in these habitats. Two females that nested on Yakushima Island, Japan, were tracked for 124 and 197 days. The small female wandered in the oceanic Pacific, and spent most of the time at 0-25 m depths regardless of day or night, implying that she foraged pelagically at the surface and shallow depths. Her mean dive durations were significantly longer at night than during the day. The large female moved into the neritic East China Sea, and spent most of the time over the continental shelf at 100-150 m depths during the day and at 0-25 m depths at night, suggesting that she alternated between diurnal benthic foraging and nocturnal resting within the depths where she could attain neutral buoyancy. Her mean dive durations were not significantly different between day and night. The increase in dive duration for both turtles coincided with a seasonal decrease in water temperature. The small female sometimes showed midwater dormancy at 0-25 m depths with a duration of 45 h that was in contrast with bottom dormancy by sea turtles inhabiting other regions. The diving behaviours observed during this study were consistent with their estimated main feeding habits, which demonstrated resource polymorphism in a marine reptile.

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Adult female loggerhead sea turtles ( Caretta caretta (L., 1758)) show a size-related behavioural and trophic dichotomy within several populations: oceanic planktivory by small females and neritic benthivory by large females. Within a Japanese population, smaller females have longer remigration intervals (the intervals between successive nesting years) than larger females. We hypothesized that this is attributed to the size-related foraging dichotomy, and calculated energy budgets and food requirements for the small and large females. Total reproductive energy costs for the small and large females were 1003 and 434 MJ, equivalent to 4043 kg of planktonic items and 275 kg of benthic items, respectively. The minimum durations necessary for accumulating the total reproductive energy for the small and large females were 430 and 26 days, equivalent to remigration intervals of 2 years and 1 year taking into account the nesting and migration durations and seasonal reproduction. Therefore, it was theoretically demonstrated that the intrapopulation variation in remigration intervals for Japanese loggerheads could be caused by the difference in their feeding habits, implying that remigration dynamics in a sea turtle population that shows a foraging dichotomy is regulated by feeding conditions not in a single habitat but in multiple habitats.