Nesting fidelity and molecular evidence for natal homing in the freshwater turtle, Graptemys kohnii

Numerous studies of sea turtle nesting ecology have revealed that females exhibit natal homing, whereby they imprint on the nesting area from which they hatch and subsequently return there to nest as adults. Because freshwater turtles comprise the majority of reptiles known to display environmental sex determination (ESD), the study of natal homing in this group may shed light on recent evolutionary models of sex allocation that are predicated on natal homing in reptiles with ESD. We examined natal homing in Graptemys kohnii, a freshwater turtle with ESD, using mitochondrial sequencing, microsatellite genotyping and mark and recapture of 290 nesting females. Females showed high fidelity to nesting areas, even after being transplanted several kilometres away. A Mantel test revealed significant genetic isolation by distance with respect to nesting locations (r=0.147; p<0.05), suggesting that related females nest in close proximity to one another. The patterns of fidelity and genotype distributions are consistent with homing at a scale that may affect population sex ratios.     

Variation in reproductive output of marine turtles

Most marine turtle species are non-annual breeders and show variation in both the number of eggs laid per clutch and the number of clutches laid in a season. Large levels of inter-annual variation in the number of nesting females have been well documented in green turtle nesting populations and may be linked to environmental conditions. Other species of marine turtle exhibit less variation in nesting numbers. This inter-specific difference is thought to be linked to trophic status. To examine whether individual reproductive output is more variable in the herbivorous green turtle (Chelonia mydas Linneaeus 1758) than the carnivorous loggerhead (Caretta caretta Linneaeus 1758), we examined the nesting of both species in Cyprus over nine seasons. Green turtles showed slower annual growth rates (0.11 cm year⁻¹ curved carapace length (CCL) and 0.27 cm year⁻¹ curved carapace width (CCW)) than loggerhead turtles (0.36 cm year⁻¹ CCL, 0.51 cm year⁻¹ CCW). CCL was highly correlated to mean clutch size in both green (R²=0.51) and loggerhead turtles (R²=0.61) and maximal clutch size of green turtles (R²=0.58). Larger females did not lay a greater number of clutches or have a shorter remigration interval than smaller females of either species. On average, the size of green turtle clutches increased and that of loggerhead turtles decreased as the season progressed. Individual green turtles, however, produced more eggs per clutch through the season to a maximum in the third or fourth clutch. In loggerhead turtles, clutches 1-4 were very similar in size but the fifth clutch was 38% smaller than the first. No individuals of either species were recorded laying more than five clutches. Green turtles may not be able to achieve their maximum reproductive output with respect to clutch size throughout the season, whereas only loggerhead turtles laying five clutches (n=5) appear to become resource depleted. Green turtles nesting in years when large numbers of nests were recorded laid a greater number of clutches than females nesting in years with lower levels of nesting.     

The Effects of Artificial Beach Nourishment on Marine Turtles: Differences between Loggerhead and Green Turtles

Marine turtle reproductive success is correlated with the stability and quality of the nesting environment. Female marine turtles show fidelity to nesting beaches, making artificial beach nourishment practices directly relevant to their recovery. We evaluated the impacts of artificial beach nourishment on Loggerhead ( Caretta caretta ) and Green turtles ( Chelonia mydas ) between artificially nourished and nonnourished beaches. We observed reduced nesting success (ratio of nesting emergences to emergences not resulting in nest deposition) for both species. This negative effect lasted for one season in Loggerheads and for at least one season in Green turtles. Physical attributes of the fill sand did not impede nesting attempts. We argue that the decrease in nesting success resulted from an altered beach profile not favorable for nest deposition, which subsequently improved in later seasons as the beach equilibrated to a more natural slope. We observed a 52.2% decrease in reproductive output (hatchlings km⁻¹ yr⁻¹) for Loggerheads one year postnourishment, with a 44.1% increase observed the two seasons postnourishment. In Green turtles, a 0.8% reduction was observed the first season postnourishment, despite a 13% increase in the nonnourished area. The reduction in reproductive output in both cases was primarily a consequence of decreased nesting success, lowering nest numbers. These results reveal stronger negative effects of beach nourishment on Loggerheads compared to Green turtles and the importance of minimizing excessive nonnesting emergences associated with artificial beach nourishment. Nourished areas also experienced more than 600% increase in the number of Loggerhead hatchlings disoriented by artificial lighting over two years postnourishment.     

Trophic status drives interannual variability in nesting numbers of marine turtles.

Large annual fluctuations are seen in breeding numbers in many populations of non-annual breeders. We examined the interannual variation in nesting numbers of populations of green (Chelonia mydas) (n = 16 populations), loggerhead (Caretta caretta) (n = 10 populations), leatherback (Dermochelys coriacea) (n = 9 populations) and hawksbill turtles (Eretmochelys imbricata) (n = 10 populations). Interannual variation was greatest in the green turtle. When comparing green and loggerhead turtles nesting in Cyprus we found that green turtles were more likely to change the interval between laying seasons and showed greater variation in the number of clutches laid in a season. We suggest that these differences are driven by the varying trophic statuses of the different species. Green turtles are herbivorous, feeding on sea grasses and macro-algae, and this primary production will be more tightly coupled with prevailing environmental conditions than the carnivorous diet of the loggerhead turtle.     

Do trade‐offs between predation pressures on females versus nests drive nest‐site choice in painted turtles?

Predation strongly influences reproductive behaviours because reproducing individuals must balance mortality risks to themselves and to their offspring. In many freshwater turtles, the nest predation risk decreases with nest distance from water, whereas the predation risk to females increases farther from water. To determine whether predation pressure influences the distance from water at which female turtles nest, we measured predation pressure on nesting females and on nests, as well as the distances of nests to water, in two populations of painted turtles. Using models, we found that female survival in both populations was high and did not vary with distance from water. Nest survival was also uncorrelated with nest distance to water, although it was significantly lower than adult survival in both populations and was only 1.2% in one population. Our results suggest that nest sites are not predictably safe from predators. Instead, turtles may hedge their bets by nesting over a wide range of distances from water because any distance is risky for nests and no distance is particularly risky for the nesting female. We suggest that other factors, such as suitable incubation conditions and/or post‐emergence hatchling survival, probably play a larger role than predation in driving nest‐site choice in painted turtles.     

Annual variation in nesting numbers of marine turtles: the effect of sea surface temperature on re-migration intervals

The assessment of marine turtle populations is complicated by large inter‐annual variations in nesting numbers. This variability has been attributed to synchrony in the re‐migration intervals (i.e., intervals between successive nesting years) of individuals. It has been hypothesized that this synchrony arises from environmental variations that affect the feeding conditions at sea. This paper presents the first direct test of this hypothesis. The analysis identifies a significant effect of sea surface temperature on the re‐migration interval in the largest Atlantic population of green turtles.     

Post-nesting migrations patterns of Green Turtles (Chelonia mydas) from the Egyptian Red Sea

In order to identify the migratory pathways and foraging grounds of post-nesting Green Turtles (Chelonia mydas) in the Red Sea, we attached satellite transmitters to four females immediately after egg deposition and tracked them between 207 and 647 days. We identified four geographically distinct post-nesting habitat areas and migration paths from Zabargad Island, Egypt which was remarkable given our small sample size. Our shortest migration was 140?km and the longest 940?km, with the migrations and post-nesting habitat encompassing the boundaries of four of the seven Red Sea countries (Egypt, Sudan, Eritrea, and Saudi Arabia). The post-nesting habitats were located in shallow coastal habitat and three consisted of near-shore archipelagos. Two turtles moved past areas of suitable post-nesting habitat that was occupied by other turtles, which suggests that these turtles may be exhibiting fidelity to certain feeding and nesting sites. Our results suggest that regional and multi-national cooperation will be needed to protect sea turtles that nest on Zabargad Island, a nesting site that is important for Egypt and other Red Sea nations.     

Nest site fidelity of leatherback turtles at Playa Grande, Costa Rica

The possibility of nest site fidelity of leatherback turtles, Dermochelys coriacea, raises concerns about the merit of translocating poorly located clutches to a safe hatchery. Specifically, if some females habitually nest in areas where the probability of egg survival is low because of tidal inundation, nest relocation may inadvertently select for turtles that make poor nest site selections. We investigate the nest site fidelity of leatherbacks by studying consecutive nest site selections by leatherbacks at Playa Grande, Costa Rica, from 1993 to 2001. Along the ocean-to-vegetation axis, consecutive nest placements were independent (G₄²=4.737, P=0.315). Regardless of previous zone choices, leatherbacks typically nested in the open beach above the high tide line and away from vegetation. Thus, concerns about translocation of poorly located nests are unfounded. Along the coastal axis, however, an independence model was rejected (G₃₆²=145.179, P<0.001). The log-linear model-based analysis we developed and applied indicated that the pattern of dependence in nest site locations is more consistent with a hypothesis of spatial proximity in which leatherbacks simply nest close to the site of the previous nest than it is with a hypothesis of intrinsic fidelity to a specific location on the beach. Our model-based approach is suitable for other settings where longitudinal nesting data are available.     

Nest Site Preference and Fidelity of Chinese Alligator(Alligator sinensis)

Oviparous female reptiles select nesting sites with optimal ecological factors that contribute to egg development. Chinese alligator(Alligator sinensis), an oviparous reptile, is a critically endangered crocodilian with temperature-dependent sex determination. Research on its nesting behavior may facilitate the protection of this species. In this study, we monitored nesting behavior over eight years. We compared selected frequency of nest sites, distance from nest site to water, height from nest site top to the water surface, distance from nest site to human activity region, and canopy density between nest sites on the island and bank. The results showed that 45 nest sites were used by female alligators over eight years and each site was selected from one to 10 times. The selected frequency of nest site occurrence on the island was higher than that on the bank(P 0.001). We observed that 88% of the individual alligators(15/17) showed different degrees of nest site fidelity. However, Chinese alligators might not always be loyal to only one nest site because of environmental changes or interspecific competition at nest sites. Our findings suggest that female alligators prefer to nest at island, which might be because of the nests on the island had a higher canopy density(P = 0.010) and were further from the human activity region(P 0.001) than those on the bank did. It would be beneficial to reduce human activities during the breeding season and protect the vegetation of Chinese alligator habitats in the future. of the beach(Ali et al., 2005; Kamel and Mrosovsky, 2005; Turkozan et al., 2012). Other species such as snakes and lizards select nest sites with more moist substrates, which significantly increases their body size at hatching(Brown and Shine, 2004; Reedy et al., 2013). Nest site preference may be maintained for one or more breeding seasons(Janzen and Morjan, 2001; Kamel et al., 2006). When female chose the same site or area during different years, it is called nest site fidelity(Switzer, 1993). This nest choice behavior may influence offspring survival(Lindeman, 1992) and nest site fidelity has been documented in numerous species such as birds(Lindberg and Sedinger, 1997), turtles(Mitrus, 2006; Walde et al., 2007), tuatara(Refsnider et al., 2010), and crocodilians(Elsey et al., 2008).Chinese alligator(Alligator sinensis) is a critically endangered freshwater crocodilian endemic to China(Thorbjarnarson and Wang, 1999). Recent investigations show that there are no more than 130 Chinese alligators     

Natural beaches confer fitness benefits to nesting marine turtles

Coastal ecosystems provide vital linkages between aquatic and terrestrial habitats and thus support extremely high levels of biodiversity. However, coastlines also contain the highest densities of human development anywhere on the planet and are favoured destinations for tourists, creating a situation where the potential for negative effects on coastal species is extremely high. I gathered data on marine turtle reproductive output from the literature to determine whether coastal development negatively influences offspring production. Female loggerhead (Caretta caretta) and green turtles (Chelonia mydas) nesting on natural beaches (as opposed to beaches with permanent development) produce significantly more hatchling turtles per nest; all else being equal, females that successfully produce more offspring will have higher fitness than conspecifics producing fewer offspring. Thus, female marine turtles nesting on natural beaches probably have higher fitness than turtles nesting on developed beaches. Consequently, populations nesting on natural beaches may be able to recover more quickly from the historic population declines that have plagued marine turtles, and some species may recover more quickly than others.     

High thermal variance in naturally incubated turtle nests produces faster offspring

The effects of climate change on populations are complex and difficult to predict, and can result in mismatches between interdependent organisms or between organisms and their environment. Reptiles with temperature-dependent sex determination may be able to compensate for potential skews in offspring sex ratio caused by climate change by selecting cooler (i.e., shadier) nest sites. Although changing nest location may prevent sex ratio skews, it may also affect thermally sensitive performance traits in offspring. I tested righting, sprinting, and swimming performance in hatchling painted turtles (Chrysemys picta), produced by female turtles from five populations across the species’ geographic range, nesting in a common-garden environment. I found that speed of hatchling performance was faster in hatchlings whose mothers originated from warmer climates, and that nests with higher mean daily variation in incubation temperature produced faster hatchlings. These results suggest that the increased temperatures predicted by climate change models could result in hatchling turtles that are faster at sprinting and swimming; however, it is not yet known how these performance measures translate into fitness.     

Nest site fidelity and clutch frequency of loggerhead turtles are better elucidated by satellite telemetry than by nocturnal tagging efforts: Implications for stock estimation

Satellite telemetry and ground-based tagging studies are complementary methods to define the spatial and temporal patterns of nesting behavior by migratory sea turtles. Estimates of site fidelity and clutch frequencies are compared for satellite telemetry versus ground truth patrols over a 6 km stretch at a southwest Florida loggerhead (Caretta caretta) rookery. Site fidelity ranged from 1.9 km to 109.1 km for all nests deposited by a female within a season. The mean site fidelity was 28.1 km for all nests, but declined to 16.9 km if omitting the first nest. Nest frequency ranged from 2 to 8 nests per season, with a modal value of 5 nests. Satellite telemetry documented a mean nest frequency of 5.4 nests per female in comparison to 2.2 nests detected by monitoring patrols. The remigrant females had higher clutch frequency, were larger in size, and had higher site fidelity compared to newly tagged females. Satellite telemetry provided improved measurements of site fidelity and reveals a need for revised fecundity estimates. If measures of clutch frequency are representative of loggerhead assemblages nesting elsewhere within the South Florida grouping, the confidence bounds on Western Atlantic loggerhead stocks are approximately 32% lower than currently accounted for annual nesting individuals.     

Estimating demographic parameters for loggerhead sea turtles using mark–recapture data and a multistate model

The survival for adult loggerhead sea turtles from a saturation tagging study on Bald Head Island, NC, USA, was estimated using a multistate model with unobservable states to relax assumptions that are violated when survival is estimated from multistate models and produce more accurate estimates of survival, recapture, and breeding transition probabilities. The influence of time, trap dependence, and low site fidelity to the study nesting beach on survival and recapture were examined. The best model given the data included an imprecise site-fidelity effect on survival, constrained the reproductive cycle to 4 years, and contained a time effect on recapture rates. The estimate of annual survival for adult females was of 0.85, producing the highest estimate in the literature for loggerhead sea turtles. Multistate models should be applied to other nesting beach data for sea turtles to improve survival estimates and in turn the ability to model and manage populations.     

The effects of rainfall on different components of seasonal fecundity in a tropical forest passerine

Seasonal fecundity is a composite metric that is determined by component parameters such as clutch size, nest survival and re‐nesting probability. Many of these component parameters are known to vary with environmental conditions, in particular rainfall prior to or during the breeding season. In some species, seasonal fecundity is positively related to rainfall, but little is known about which component parameters of seasonal fecundity respond most strongly to rainfall. We used intensive nest monitoring of a multi‐brooded tropical forest passerine, the Montserrat Oriole Icterus oberi, to examine the effects of rainfall during the pre‐breeding season on component parameters of annual fecundity. We monitored all nests of a total of 42 pairs over 5 years in which rainfall varied substantially. We then related clutch size, nest survival, onset and length of the breeding season, re‐nesting probability and re‐nesting interval to pre‐breeding season rainfall using generalized linear mixed models that accounted for random variation across sites and individual pairs, and incorporated other variables known to affect the response. Higher pre‐breeding season rainfall led to an increase in clutch size and a decrease in re‐nesting interval, but nest survival, re‐nesting probability and length of the breeding season were not affected by variation in rainfall. The onset of the breeding season was delayed in very dry years. We conclude that higher rainfall is likely to increase food availability and thus body condition of female Montserrat Orioles, leading to an increase in fecundity due to larger clutch sizes.     

International migrations of South American sea turtles (Cheloniidae and Dermochelidae)

Three species of sea turtles (the leatherback, Dermochelys coriacea; the green turtle, Chelonia mydas; and the olive ridley, Lepidochelys olivacea) nest abundantly in the Guianas, especially on the beaches adjacent to the mouth of the Marowijne River. Tagging demonstrated that green turtles nesting in Surinam are recruited from feeding grounds in or near the State of Ceará, Brazil, while olive ridleys, after nesting in Surinam, spread out over 3800 km of the coast of northern South America. A single tagged leatherback was recovered in Ghana.     

Latitudinal trends in within-year reoccupation of nest boxes and their implications

Multiple brooding can substantially increase the annual reproductive output of birds, and the propensity for multiple brooding can vary geographically. Thus, studies attempting to understand the evolution of geographic variation in nesting success need to account for variation in re‐nesting potential. However, direct assessment of rates of multiple brooding requires individually recognizable breeding adults, which are not generally available. We explore the possibility of comparing relative indices of multiple broodedness across a latitudinal gradient from studies of un‐banded birds locally restricted to nest boxes. We analyzed nest box reoccupation by a multiple‐brooding species, the eastern bluebird Sialia sialis, reported by volunteers in a citizen‐participation project (1998–2002) in which nest boxes were monitored throughout much of the breeding range of the bluebirds. We found nest boxes in the southern portion of the bluebird range (30° latitude) had, on average 17–33% higher likelihood of repeated egg‐laying, brooding, and successful fledging events than boxes in the north (48° latitude). Latitudinal variation in the reoccupation of nest boxes may indicate that either (1) the number of broods per female varies with latitude, (2) female breeding dispersal/site fidelity varies with latitude, (3) the density, distribution, and/or availability of suitable nest sites varies with latitude, or (4) observer bias varies with latitude. Various lines of evidence suggest that nest re‐occupancy is a useful index of latitudinal variation in re‐nesting. During the time‐frame of second attempts, first‐time box occupancy was as likely as second occupancy and approximately 45% more likely in the south than north, suggesting that, despite considerable breeding dispersal, observed trends in box reoccupation conservatively reflect latitudinal trends in the number of nest attempts/broods per female. Furthermore, despite a compressed nesting cycle in the north (shorter incubation and re‐nesting interval), the shorter duration of the breeding season in the north restricted the potential number of broods. Studies of banded birds are necessary to confirm the behavior underlying the latitudinal trends in box reoccupation.     

Nesting Habitat Creation Enhances Recruitment in a Predator‐Free Environment: Malaclemys Nesting at the Paul S. Sarbanes Ecosystem Restoration Project

Aquatic turtles worldwide are plagued with habitat loss due to development and shoreline alteration that destroys the terrestrial–aquatic linkage which they must cross to reproduce successfully. Furthermore, nesting habitat loss can concentrate nesting, increasing nest predator efficiency. We describe how the Paul S. Sarbanes Ecosystem Restoration Project at Poplar Island created nesting habitat for Malaclemys terrapin (Diamondback Terrapin), and document nesting success in response to construction progress and the absence of raccoons and foxes, the primary nest predators. We monitored terrapin nests throughout the nesting seasons from 2002 to 2011 to determine overall and within‐nest survivorship. Female terrapins began nesting on the restoration project within 1 year but planned construction during the study eliminated some nesting areas and opened previously inaccessible areas. Overall, nest survivorship was considerably higher than mainland nesting areas due to the absence of raccoons and foxes on the island and within‐nest survivorship was similar. Egg size, hatchling size, and the frequency of shell scute anomalies were similar to other terrapin populations, suggesting normal developmental conditions on the island. We documented annual variation in hatchling size that correlated negatively with mean air temperature during the incubation season. Our results indicate that restored or created isolated island habitat can be located rapidly by terrapins and can become an important source of recruitment in regions where nesting habitat is limited and predation is high. Poplar Island illustrates how habitat loss and restoration can affect turtle populations by revealing the changes in nesting patterns and success in newly created, predator‐free habitat.     

A Comparison of the Seasonal Movements of Tiger Sharks and Green Turtles Provides Insight into Their Predator-Prey Relationship

During the reproductive season, sea turtles use a restricted area in the vicinity of their nesting beaches, making them vulnerable to predation. At Raine Island (Australia), the highest density green turtle Chelonia mydas rookery in the world, tiger sharks Galeocerdo cuvier have been observed to feed on green turtles, and it has been suggested that they may specialise on such air-breathing prey. However there is little information with which to examine this hypothesis. We compared the spatial and temporal components of movement behaviour of these two potentially interacting species in order to provide insight into the predator-prey relationship. Specifically, we tested the hypothesis that tiger shark movements are more concentrated at Raine Island during the green turtle nesting season than outside the turtle nesting season when turtles are not concentrated at Raine Island. Turtles showed area-restricted search behaviour around Raine Island for ∼3–4 months during the nesting period (November–February). This was followed by direct movement (transit) to putative foraging grounds mostly in the Torres Straight where they switched to area-restricted search mode again, and remained resident for the remainder of the deployment (53–304 days). In contrast, tiger sharks displayed high spatial and temporal variation in movement behaviour which was not closely linked to the movement behaviour of green turtles or recognised turtle foraging grounds. On average, tiger sharks were concentrated around Raine Island throughout the year. While information on diet is required to determine whether tiger sharks are turtle specialists our results support the hypothesis that they target this predictable and plentiful prey during turtle nesting season, but they might not focus on this less predictable food source outside the nesting season.     

Do female green turtles (Chelonia mydas) exhibit reproductive seasonality in a year-round nesting rookery?

In circumglobal populations of sea turtles, little nesting activity occurs during cold seasons when the sand temperature on a nesting beach is inappropriate for successful egg incubation. However, it is not known whether reproductive seasonality also occurs in year-round nesting rookeries where the sand temperature may be suitable throughout the year. Therefore, we examined the hypothesis that female turtles exhibit reproductive seasonality even in a year-round nesting population. To determine whether the time of nesting of individuals fluctuates between successive nesting seasons, a year-round nesting rookery of green turtles Chelonia mydas at Huyong Island, Thailand, was patrolled on foot every night for 3190 consecutive days, and nesting females were identified using microchips and metal tags. The date of first nesting within a season for an individual (nesting date) was considered the nesting season of each individual. We identified 94 females and 463 nests (including 47 unidentified nests) during the survey. Nesting dates were distributed throughout the year. Nineteen turtles remigrated to the nesting beach, and the nesting dates of the remigrated females occurred in the same seasons. This indicates that females have a fixed individual nesting season even in a year-round nesting rookery. The year-round nesting of green turtles may be attributed to a wide variation in the nesting seasons of individuals. Satellite tracking revealed that the mean travelling duration of post-nesting migrations in this population was 13.5±6.1 days. The proximity of locations for reproduction and feeding may also play an important role in sustaining year-round nesting in this population.     

Potential consequences of angling on nest-site fidelity in largemouth bass

Breeding site fidelity has evolved in many vertebrate taxa, suggesting both that site selection has an important influence on fitness potential and that the decision to reuse a nesting site is related to the individual’s prior nesting success at that location. For a species that provides parental care, such as the Largemouth Bass Micropterus salmoides, catch-and-release angling impacts individual nesting success and fitness through physiological disturbance and by removing the nest-guarding male from its brood, thereby allowing temporary access to eggs and hatchlings by brood predators. To assess the impact of catch-and-release angling on nest site fidelity, we compared the consequences of angling on individually marked (i.e., with passive integrated transponders) nest-guarding male Largemouth Bass in Ontario. An extremely high degree of nest site fidelity in year two was observed for males that were angled only once during year one (87% within 10 m of the previous year’s nest), 96.7% of which remained on the nest and completed parental care activities. There was significantly lower fidelity in year two, however, for males that were angled multiple times during year one (27% within 10m of the previous year’s nest), only 5.6% of which remained on the nest and completed parental care activities. This observed difference suggests that angling nesting bass may cause them to avoid previously used nest sites and instead search for alternative sites during future reproductive seasons. This human-induced impact on nest site choice may impact the future reproductive success of those Largemouth Bass.