Inter-seasonal maintenance of individual nest site preferences in hawksbill sea turtles

Within a single population of hawksbill sea turtles (Eretmochelys imbricata), we found a behavioral polymorphism for maternal nest site choice with respect to beach microhabitat characteristics. Some females preferred to nest in littoral forest and in places with overstory vegetation cover, and others preferred to nest in more open, deforested areas. Nest site choice was consistent within and between nesting seasons two years apart. This was not a result of females simply returning to the same location along the shoreline; beach sections used by individual turtles varied between seasons. Nest site choice was not influenced by changes in beach environment (e.g., beach width and foliage cover) or by changes in females' reproductive output (e.g., clutch size), suggesting that fidelity to particular microhabitats is a major determinant of the observed nesting patterns. Because hawksbills exhibit temperature-dependent sex determination, if the behavioral polymorphism in nest site choice has a genetic basis, as is plausible, then this would have implications for sex ratio evolution and offspring survival. By taking an individual-based approach to the study of maternal behavior we reveal previously overlooked individual variation and hope to provide some impetus for more detailed studies of nest site choice.     

Intra-Mediterranean migrations of loggerhead sea turtles (Caretta caretta) monitored by satellite telemetry

The movements of four Mediterranean loggerhead sea turtles (Caretta caretta; three females, one male) were tracked via satellite telemetry for between 108 and 457 days. Total length of the routes traveled by the turtles varied between 2554 and 7098 km, and the average travel rate was 1.2 km h^-1. Long-distance movement between the western and eastern Mediterranean basins followed a seasonal pattern and seemed to be triggered by temperature and food availability. In the autumn/winter months turtles moved from west to east in search of warmer waters, and returned to the western basin in spring, where food resources are generally more plentiful. Three (two females, one male) of the four turtles migrated eastward through the Straits of Messina, which is characterized by high fishing pressure and intense boat traffic. Information about turtle migration patterns and routes will serve to plan effective conservation strategies.     

Female reproductive success and nest predation in polyterritorial wood warblers (Phylloscopus sibilatrix)

Summary Polyterritorial polygyny in passerine birds, i.e., when already-paired males attract females in secondary territories, has been explained by male deception. Secondary females are assumed to make a poor choice, because they lack information about males' pairing status. To date, the deception hypothesis has focused mainly on the hole-nesting pied flycatcher. In this study of polyterritorial wood warblers, an open-nesting species, secondary females did not differ in number of fledged young compared with females of monogamous males, and they fledged even more young than primary females. Nest predation influenced the allocation of paternal care. Almost half of the secondary females had exclusive male assistance because nests of primary females were preyed upon.     

Self-monitoring of reproductive success: nest switching in the redlip blenny (Pisces: Blenniidae)

Summary In the redlip blenny Ophioblennius atlanticus, commonly found on Caribbean reefs, eggs are laid in a male's nest and are guarded by the male until they hatch as planktonic larvae. Male reproductive success, defined as the number of egg batches hatched per reproductive period, was positively correlated with male size and with the inner surface area of the male's nest. Male blennies often switch nests between reproductive periods. Most nest switching (83%) occurred following periods of low reproductive success. Moreover, a male having low reproductive success was more likely to switch if the inner surface area of his nest was small. Most nest switching (72–80%) was to a nest of larger surface area than the previous nest. In 72% of all switches, males had higher reproductive success in the period following the switch than in the one preceding it. The results suggest that male redlip blennies monitor their current reproductive success and the characteristics of their current nests, switch to better nests if their reproductive success is low, and typically benefit from the decision to do so. These capabilities have seldom been demonstrated in any phylum.     

Testing the navigational abilities of ocean migrants: displacement experiments on green sea turtles (Chelonia mydas)

Like many animals migrating through the oceans, sea turtles face difficult navigational tasks when they have to reach distant, specific sites. The paradigmatic case of Brazilian green turtles (Chelonia mydas), which nest on the tiny Ascension Island in the middle of the Atlantic Ocean, has often been the subject of hypotheses concerning their navigational mechanisms. To investigate their nature, we displaced 18 females from Ascension and tracked them by satellite after release from eight different points in the ocean, 60-450 km away from the island. Four turtles moved to Brazil soon after the release, 4 moved in various directions before heading to Brazil, and 10 reached the island. All the successful trips, bar 1, were winding but ended with a final straight segment of variable length, as if the turtles were searching for a sensory contact with the island which they obtained at various distances. The approach to Ascension mostly occurred from the direction opposite to the trade wind, suggesting a navigational role of wind-borne information originating from the island.     

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.     

Magnetic orientation by hatchling loggerhead sea turtles (Caretta caretta) from the Gulf of Mexico

Loggerhead sea turtle hatchlings emerge from nests on either the east or west coast of the South Florida peninsula and then migrate offshore in opposite directions. Under laboratory conditions, magnetic cues induce east coast hatchlings to swim in directions that promote their transport by oceanic surface currents, such as the North Atlantic gyre. However, the surface currents used by west coast hatchlings are unknown. We examined the responses of west (Sarasota) hatchlings to magnetic cues in the Gulf of Mexico, the Florida Straits, and the Gulf Stream to determine their (1) likely migratory routes (2) orientation where currents lead into the Atlantic Ocean, and (3) orientation adjacent to Florida’s east coast. The results suggest that migration inside Gulf waters may be circuitous, that the turtles respond appropriately to enter Atlantic waters, and that orientation along Florida’s east coast probably promotes transport by the Gulf Stream into the North Atlantic gyre.     

Influence of emergence success on the annual reproductive output of leatherback turtles

Reproductive output of leatherback turtles (Dermochelys coriacea) is affected by the stochastic nature of emergence success. Average emergence success of nests at Playa Grande, Costa Rica was 0.38 ± 0.27. Incubation temperature affected development of leatherback turtle eggs and emergence of hatchlings from the nest. We found that high temperatures reduced hatching success and emergence rate and increased embryonic mortality both early and late during incubation at Playa Grande. There was a temporal effect on emergence success that resulted in more hatchlings being produced at the beginning of the season, because of higher emergence success, than toward the end. Likewise, production of hatchlings varied from year to year. The average annual reproductive output was 252 ± 141 hatchlings per female. The 2005–2006 nesting season had the highest emergence success and produced the greatest number of hatchlings per female compared to the 2004–2005 (+120%) and 2006–2007 (+41%) seasons. However, average clutch size (62 ± 10) and clutch frequency (9.45 ± 1.63), were not different among years. Turtles that had nested a high number of years exhibited greater clutch frequency and arrived earlier to nest than turtles that had nested in fewer numbers of years. Nesting when environmental conditions favor high developmental success and emergence rate may constitute an advantageous reproductive strategy.     

Variation in nesting patterns affecting nest temperatures in two populations of painted turtles (<Emphasis Type="Italic">Chrysemys picta</Emphasis>) with temperature-dependent sex determination

Mechanisms maintaining sex ratios in populations with temperature-dependent sex determination (TSD) remain elusive. Although geographic variation in embryonic sex determination (i.e., pivotal temperature) has been widely investigated in reptiles exhibiting TSD, no previous studies have directly addressed geographic variation in maternal behavior affecting nest thermal conditions. I evaluated patterns of nest-site selection and its effects on thermal and hydric nest conditions for a population of painted turtles ( Chrysemys picta bellii) exhibiting TSD in New Mexico. These results are compared to data collected from a well-studied, conspecific population experiencing relatively cooler climatic conditions in Illinois. Since canopy vegetation cover reduces nest temperatures in Illinois, I expected females in New Mexico to nest under high amounts of canopy vegetation cover. However, females from New Mexico placed nests under significantly less canopy vegetation cover, but closer to standing water, than did females from Illinois. Experimental nests in New Mexico demonstrated that increased canopy vegetation cover and soil moisture reduced nest temperatures. By nesting close to standing water rather than under canopy vegetation cover, females in New Mexico nested in habitats more closely associated with maximizing moisture around nests rather than reducing nest temperatures through shading. Mean July nest temperatures were similar for both populations. Since nest hydric conditions affect hatching success and hatchling size in C. picta, nesting patterns in New Mexico may primarily reflect selection for microhabitats affecting offspring survivorship or size.     

Fieldfare (Turdus pilaris) breeding success in relation to colony size,nest position and association with Merlins (Falco columbarius)

Summary Clutch size, nestling production and breeding success were studied in colonial Fieldfares (Turdus pilaris) in a subalpine birch forest during ten breeding seasons. Reproductive success was highest for central pairs in large colonies; such pairs benefited most from communal defence against nest predators. Fieldfares and Merlins (Falco columbarius) usually bred in association. Fieldfares breeding away from Merlins had lower breeding success than pairs associated with Merlins, which also benefited by reduced nest predation. Fieldfares apparently chose to nest near Merlins, which had already laid eggs when the thrushes started nest-building.     

Long-term behavior at foraging sites of adult female loggerhead sea turtles (Caretta caretta) from three Florida rookeries

We used satellite telemetry to study behavior at foraging sites of 40 adult female loggerhead sea turtles (Caretta caretta) from three Florida (USA) rookeries. Foraging sites were located in four countries (USA, Mexico, the Bahamas, and Cuba). We were able to determine home range for 32 of the loggerheads. One turtle moved through several temporary residence areas, but the rest had a primary residence area in which they spent all or most of their time (usually >11 months per year). Twenty-four had a primary residence area that was <500 km² (mean = 191). Seven had a primary residence area that was ≥500 km² (range = 573–1,907). Primary residence areas were mostly restricted to depths <100 m. Loggerheads appeared to favor areas with larger-grained sediment (gravel and rock) over areas with smaller-grained sediment (mud). Short-term departures from primary residence areas were either looping excursions, typically involving 1–2 weeks of continuous travel, or movement to a secondary residence area where turtles spent 25–45 days before returning to their primary residence area. Ten turtles had a secondary residence area, and six used it as an overwintering site. For those six turtles, the primary residence area was in shallow water (<17 m) in the northern half of the Gulf of Mexico (GOM), and overwintering sites were farther offshore or farther south. We documented long winter dive times (>4 h) for the first time in the GOM. Characterizing behaviors at foraging sites helps inform and assess loggerhead recovery efforts.     

Genetic structuring of immature loggerhead sea turtles (Caretta caretta) in the Mediterranean Sea reflects water circulation patterns

The analysis of mitochondrial DNA in loggerhead sea turtles (Caretta caretta) from eight foraging grounds in the Mediterranean and the adjoining Atlantic revealed deep genetic structuring within the western Mediterranean. As a consequence, the foraging grounds off the North-African coast and the Gimnesies Islands are shown to be inhabited mainly by turtles of the Atlantic stocks, whereas the foraging grounds off the European shore of the western Mediterranean are shown to be inhabited mainly by turtles from the eastern Mediterranean rookeries. This structuring is explained by the pattern of sea surface currents and water masses and suggests that immature loggerhead sea turtles entering the western Mediterranean from the Atlantic and the eastern Mediterranean remain linked to particular water masses, with a limited exchange of turtles between water masses. As the north of the western Mediterranean comprises mostly individuals from the highly endangered eastern Mediterranean rookeries, conservation plans should make it a priority to reduce the mortality caused by incidental by-catch in these areas.     

Multiple paternity, sperm storage, and reproductive success of female and male painted turtles (Chrysemys picta) in nature

When females receive no direct benefits from multiple matings, concurrent multiple paternity is often explained by indirect genetic benefits to offspring. To examine such possibilities, we analyzed genetic paternity for 1,272 hatchlings, representing 227 clutches, from a nesting population of painted turtles (Chrysemys picta) on the Mississippi River. Goals were to quantify the incidence and distribution of concurrent multiple paternity across clutches, examine temporal patterns of sperm storage by females, and deduce the extent to which indirect benefits result from polyandrous female behaviors. Blood samples from adult males also allowed us to genetically identify the sires of surveyed clutches and to assess phenotypic variation associated with male fitness. From the genetic data, female and male reproductive success were deduced and then interpreted together with field data to evaluate possible effects of female mating behaviors and sire identity on offspring fitness. We document that more than 30% of the clutches were likely fathered by multiple males, and that presence of multiple paternity was positively correlated with clutch size. Furthermore, the data indicate that the second male to mate typically had high paternity precedence over the first.     

Conservation implications of density-dependent predation by ghost crabs on hatchling sea turtles running the gauntlet to the sea

Protecting eggs from predators is common practice in sea turtle conservation, but routine protection of hatchlings is not. Of 42 loggerhead hatchlings observed emerging from 10 nests on undeveloped Onslow Beach, North Carolina, 24 % were preyed on by ghost crabs. In experimental trials, ghost crabs similarly threatened and captured neonate freshwater sliders, supporting their substitution as proxy for threatened and endangered sea turtle hatchlings in field experiments testing density dependence. Exploiting natural long-shore variation in ghost crab density, we show that a 2.6-fold higher ghost crab density resulted in 5 times more nocturnal threat encounters with sliders and 3.4 times more slider captures. Sliders released in simulated group emergences experienced lower per capita capture risk by ghost crabs than solitary sliders, implying predator dilution. Non-independence of egg and hatchling depredation motivates consideration of merging sea turtle egg and hatchling stages when modeling and managing food web interactions.     

Temporal, spatial, and body size effects on growth rates of loggerhead sea turtles (Caretta caretta) in the Northwest Atlantic

In response to a call from the US National Research Council for research programs to combine their data to improve sea turtle population assessments, we analyzed somatic growth data for Northwest Atlantic (NWA) loggerhead sea turtles (Caretta caretta) from 10 research programs. We assessed growth dynamics over wide ranges of geography (9–33°N latitude), time (1978–2012), and body size (35.4–103.3 cm carapace length). Generalized additive models revealed significant spatial and temporal variation in growth rates and a significant decline in growth rates with increasing body size. Growth was more rapid in waters south of the USA (<24°N) than in USA waters. Growth dynamics in southern waters in the NWA need more study because sample size was small. Within USA waters, the significant spatial effect in growth rates of immature loggerheads did not exhibit a consistent latitudinal trend. Growth rates declined significantly from 1997 through 2007 and then leveled off or increased. During this same interval, annual nest counts in Florida declined by 43 % (Witherington et al. in Ecol Appl 19:30–54, 2009) before rebounding. Whether these simultaneous declines reflect responses in productivity to a common environmental change should be explored to determine whether somatic growth rates can help interpret population trends based on annual counts of nests or nesting females. Because of the significant spatial and temporal variation in growth rates, population models of NWA loggerheads should avoid employing growth data from restricted spatial or temporal coverage to calculate demographic metrics such as age at sexual maturity.     

Satellite tracking of migrating loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) displaced in the open sea

Loggerhead turtles (Caretta caretta) are known to migrate towards fixed, individually-specific residential feeding grounds. To study their spatial behaviour and their navigational ability, five loggerheads nesting in South Africa were captured when about to start their postnesting migration and tracked by satellite after having been displaced from their usual migratory route. The first turtle, released south of Madagascar about 1,148 km from the capture site, moved west up to mainland Africa and then reached her feeding grounds by following the coast. A second turtle, released farther away (2,140 km) close to La Réunion Island, stopped for some time on the Madagascar east coast, then turned southwards to round the island and regain the African mainland in the northwest, without however allowing us to establish the location of her residential grounds. Three other turtles were released off the Tanzanian coast, 2,193 km north of their nesting area, at the northern edge of the distribution of the feeding grounds along the African coast. All of them headed north, and one turtle found her residential grounds located north of the release site. The other two females started long-distance oceanic wanderings in which they crossed nearly the entire Indian Ocean, apparently being transported by the sea currents of the region. We conclude that adult loggerhead turtles are apparently unable to compensate for the displacement and can return to a pelagic life style characteristic of juvenile turtles. These findings suggest that South African loggerheads rely on simple orientation mechanisms, such as the use of the coastline, as a guide, and compass orientation, possibly integrated by spatiotemporal programmes and/or acquired maps of familiar sites.Communicated by R. Cattaneo-Vietti, Genova     

High-use oceanic areas for Atlantic leatherback sea turtles (Dermochelys coriacea) as identified using satellite telemetered location and dive information

The movements and behavior of nine female leatherback sea turtles, Dermochelys coriacea (L.) were monitored for up to 370 days from their nesting beaches on the Caribbean island of Trinidad between 1995 and 2004 using satellite-linked time and depth recorders. During the inter-nesting period (typically March–July) turtles ranged widely, but frequented the area around Galera Point on the NE corner of Trinidad. Diving depths were typically <51 m. Upon leaving Trinidad, the three longest tracked turtles moved to higher latitude foraging areas, NE of the Flemish Cap; along the continental shelf of the Iberian peninsula to the Bay of Biscay; and along the N. Atlantic subtropical front, where they remained until the end of November. Dives were initially deep (100–300 m) and long (>26 min) as the turtles left the Caribbean, but became very shallow (>50 m) and short at high latitudes. Between mid-October and mid-November, the turtles left high latitudes for a presumed foraging area in the Mauritania upwelling where they resided until their tracking records ended. Diving remained relatively shallow. It is proposed that movements of these turtles from one foraging area to another are driven by the opportunity to forage in areas of distinct oceanic structure which serve to concentrate their gelatinous prey (e.g., salps, Scyphomedusae, Siphonophora) either at or below the surface.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Effects of nest site concealment on hatching success,reproductive success,and paternal behavior of the threespine stickleback,Gasterosteus aculeatus

Summary Male sticklebacks (Gasterosteus aculeatus) actively competed for a limited number of clay flowerpots as nest sites in circular wading pools. Males nesting in pots spawned earlier and more often, had higher mean and lower variance for hatching success (number of fry per gram of estimated clutch weight), and suffered fewer stolen fertilizations, nest raids, and territorial encounters than did males nesting outside pots, and in pools without pots (Table 2). Males nesting in pots had the more even temporal distribution of fanning bouts and interfanning intervals; the durations of fanning bouts and interfanning intervals were less variable for these males (Table 2).In replicate aquaria, one male nested on each side of a glass partition and both males spawned approximately simultaneously. Flowerpots were provided to one, both, or neither male. Males with pots had higher mean hatching success, fewer territorial encounters, and the more even temporal distribution of fanning bouts and interfanning intervals (Table 4).Thus it appears that nest site concealment, in the form of clay flowerpots, affected male hatching success, reproductive success, and parental behavior, even when physical contact between males was prevented. These data suggest that a causal relationship may exist between the paternal fanning regime and the subsequent hatching success. Furthermore, these data suggest that males which nest in high concealment increase their reproductive success, and that females which spawn with these males reduce the variance of their hatching success as well as increase their mean hatching success.     

Mitigating the effects of climate change on the nests of sea turtles with artificial irrigation

For sea turtles, like many oviparous species, increasing temperatures during development threaten to increase embryonic mortality, alter offspring quality, and potentially create suboptimal primary sex ratios. Various methods are being implemented to mitigate the effects of climate change on reproductive success, but these methods, such as breeding programs, translocations, and shading, are often invasive and expensive. Irrigation is an alternative strategy for cooling nests that, depending on location, can be implemented relatively quickly and cheaply. However, multiple factors, including ambient conditions, nest substrate, and species characteristics, can influence irrigation success. Additionally, irrigation can vary in duration, frequency, and the volume of water applied to nests, which influences the cooling achieved and embryonic survival. Thus, it is critical to understand how to maximize cooling and manage risks before implementing irrigation as a nest-cooling strategy. We reviewed the literature on nest irrigation to examine whether artificial irrigation is feasible as a population management tool. Key factors that affected cooling were the volume of water applied and the frequency of applications. Embryonic responses varied with species, ambient conditions, and the timing of irrigation during development. Nest inundation was the key risk to a successful irrigation regime. Future irrigation regimes must identify clear targets, either primary or adult sex ratios, that maximize population viability. Monitoring population responses and adjusting the irrigation regime in response to population characteristics will be critical. Most studies reported on the manipulation of only one or two variables, further research is required to understand how altering multiple factors in an irrigation regime influences the cooling achieved and embryonic responses.