Breeding biology of the green turtle, Chelonia mydas (Reptilia: Cheloniidae), on Wan-An Island, PengHu archipelago. II. Nest site selection

Nest site selection of the green turtles on Wan-An Island in the summer of 1996 was determined. Turtles (Chelonia mydas) laid on average one clutch for every three emergences. Even though the total track length was 115 m on average, individual lengths varied considerably depending on the nesting beach where the turtles emerged. Limited accessibility, i.e. adequate distance from the nearest village and a well-protected environment, make beaches A and D suitable nesting beaches for green turtles on Wan-An Island. Both total track and nesting track apexes were found clustered in the interface zone, and turtles preferred to reach the vegetation zone once they emerged from the sea. It is suggested that the turtles on Wan-An Island exhibit nest site selection behavior. Based on these results and the high nest site fidelity to their first nesting beach, conservation recommendations are proposed to the county and central governments for the preservation of nesting beaches in their natural state, by prohibiting illegal sand mining and properly controlling turtle watch groups on Wan-An Island. Received: 21 November 1997 / Accepted: 24 December 1998     

Nest-Site Selection in Individual Loggerhead Turtles and Consequences for Doomed-Egg Relocation

Abstract:  Relocation of eggs is a common strategy for conservation of declining reptilian populations around the world. If individuals exhibit consistency in their nest-site selection and if nest-site selection is a heritable trait, relocating eggs deposited in vulnerable locations may impose artificial selection that would maintain traits favoring unsuccessful nest-site selection. Conversely, if most individuals scatter their nesting effort and individuals that consistently select unsuccessful nest sites are uncommon, then artificial selection would be less of a concern. During the 2005 nesting season of loggerhead turtles ( Caretta caretta ) at Mon Repos beach, Queensland, Australia, we measured the perpendicular distance from the original nest site to a stationary dune baseline for in situ (unrelocated) and relocated clutches of eggs. We observed the fate of in situ clutches and predicted what would have been the fate of relocated clutches if they had not been moved by mapping tidal inundation and storm erosion lines. In 2005 turtles deposited an average of 3.84 nests and did not consistently select nest sites at particular distances from the stationary dune baseline. Selection of unsuccessful nest sites was distributed across the nesting population; 80.3% of the turtles selected at least one unsuccessful nest site and when previous breeding seasons were included, 97% selected at least one unsuccessful nest site. Females with nesting experience selected more successful nest sites than females with little or no experience. Relocating eggs vulnerable to tidal inundation and erosion saves the progeny from a large percentage of the population and the progeny from individuals who may in subsequent years nest successfully. Our results suggest that doomed-egg relocation does not substantially distort the gene pool in the eastern Australian loggerhead stock and should not be abandoned as a strategy for the conservation of marine turtle populations.     

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.     

The effect of incubation temperature on hatchling quality in the olive ridley turtle, Lepidochelys olivacea, from Alas Purwo National Park, East Java, Indonesia: implications for hatchery management

Nest protection through egg relocation from natural nests into protected hatcheries is a common practice used at rookeries around the world to increase hatchling recruitment into sea turtle populations. However, rarely have the impacts of this practice on hatchling recruitment and quality been assessed. This study investigated the influences of the thermal nest environment of olive ridley turtles Lepidochelys olivacea on emergence success and quality of hatchlings of hatchery nests in Alas Purwo National Park, East Java, Indonesia (2009 and 2010 nesting seasons). Nest temperatures above 34?°C for at least 3 consecutive days during incubation in the hatchery resulted in decreases in emergence success and locomotor performance of hatchlings. The use of the hatchery is recommended due to extremely high predation rate of nests left on the beach; however, altering hatchery management practice by spacing nests one meter apart and providing shade should improve hatchery outcomes now and into the future.     

Tracking hawksbill sea turtles (<Emphasis Type="Italic">Eretmochelys imbricata</Emphasis>) during inter-nesting intervals around Barbados

Hawksbill sea turtles (Eretmochelys imbricata) nesting in Barbados (Needham’s Point, 13° 04′ 41.33′′ N, 59° 36′ 32.69′′W) were outfitted with GPS dataloggers over three breeding seasons (2008–2010) to track movement during inter-nesting intervals. Most females established spatially restricted resident areas up current and within 7 km of the nesting beach where they spent the majority of the inter-nesting interval. Females nesting earlier in the season settled on shallower sites. Only experienced remigrant turtles occupied the most distant resident areas. Females tracked for multiple inter-nesting intervals exhibited site fidelity, but the area contracted and the activity of females decreased with each successive interval. Hawksbills may trade off site characteristics with distance from the nesting beach and reduce activity over the course of the breeding season to optimise energy reserves during inter-nesting intervals.     

Deforestation: risk of sex ratio distortion in hawksbill sea turtles.

Phenotypic sex in sea turtles is determined by nest incubation temperatures, with warmer temperatures producing females and cooler temperatures producing males. The common finding of highly skewed female-biased hatchling sex ratios in sea turtle populations could have serious repercussions for the long-term survival of these species and prompted us to examine the thermal profile of a relatively pristine hawksbill nesting beach in Guadeloupe, French West Indies. Data loggers placed at nest depth revealed that temperatures in the forested areas were significantly cooler than temperatures in the more open, deforested areas. Using these temperatures as a predictor of sex ratio, we were able to assess the relative contributions of the different beach zones to the primary sex ratio: significantly more males were likely to be produced in the forested areas. Coastal forests are therefore important male-producing areas for the hawksbill sea turtle, and this has urgent conservation implications. On Guadeloupe, as on many Caribbean islands, deforestation rates are high and show few signs of slowing, as there is continual pressure to develop beachfront areas. The destruction of coastal forest could have serious consequences both in terms of local nesting behavior and of regional demography through the effects on population sex ratios. Human alterations to nesting habitat in other reptile taxa have been shown to modify the thermal properties of nest sites in ways that can disrupt their ecology by allowing parasite transmission, increasing vulnerability to climate change, or rendering existing habitat unsuitable.     

The impact of behavioral and physiological maternal effects on offspring sex ratio in the common snapping turtle,<Emphasis Type="Italic"> Chelydra serpentina</Emphasis>

Theory suggests that maternal effects are especially important in organisms with environmentally-sensitive sex-determining mechanisms. However, there is no substantive body of empirical evidence to confirm this conjecture. We integrated field and laboratory studies to jointly evaluate the significance of behavioral (nest-site choice) and physiological (yolk hormone allocation) maternal effects on offspring sex ratio in the common snapping turtle (Chelydra serpentina), a species with temperature-dependent sex determination (TSD). Of the 16 microhabitat variables measured, only three (south, east, and total overstory vegetation cover) were significantly correlated with nest temperature: cooler nests were located under more vegetation cover. In turn, these microhabitat predictors of nest temperature, and nest temperature itself, may influence nest sex ratio: shadier, cooler nests were more likely to produce a higher proportion of male offspring than less shady, warmer nests. Analysis of eggs from these same nests incubated in a common garden design in the laboratory revealed that clutch sex ratio was unaffected by levels of yolk estradiol, yolk testosterone, or their interaction. Examination of both behavioral and physiological maternal effects revealed no concordant impact on offspring sex ratio. However, eggs from nests that produced male-biased sex ratios in the field yielded higher proportions of males under constant-temperature conditions in the laboratory. Our study confirms the importance of behavioral maternal effects in nature on offspring sex ratios in species with TSD, while also revealing the potential presence of a predisposition for sex-ratio production underlying TSD in this system.Communicated by S. Krackow     

Dune vegetation fertilization by nesting sea turtles

Sea turtle nesting presents a potential pathway to subsidize nutrient-poor dune ecosystems, which provide the nesting habitat for sea turtles. To assess whether this positive feedback between dune plants and turtle nests exists, we measured N concentration and delta15N values in dune soils, leaves from a common dune plant (sea oats [Uniola paniculata]), and addled eggs of loggerhead (Caretta caretta) and green turtles (Chelonia mydas) across a nesting gradient (200-1050 nests/km) along a 40.5-km stretch of beach in east central Florida, USA. The delta15N levels were higher in loggerhead than green turtle eggs, denoting the higher trophic level of loggerhead turtles. Soil N concentration and delta15N values were both positively correlated to turtle nest density. Sea oat leaf tissue delta15N was also positively correlated to nest density, indicating an increased use of augmented marine-based nutrient sources. Foliar N concentration was correlated with delta15N, suggesting that increased nutrient availability from this biogenic vector may enhance the vigor of dune vegetation, promoting dune stabilization and preserving sea turtle nesting habitat.     

Song rate as a signal for nest site quality in blackcaps (Sylvia atricapilla)

In this study we examine male song output as a measure of nest site quality in blackcaps (Sylvia atricapilla). Song rate, breeding success, predation on nests and reaction to playbacks were investigated in individual males. Habitat features determining nest site and song post quality in terms of vegetation cover were compared between successful nests and nests that had suffered predation. We then related song rate of unmated males to habitat factors in territories and nesting sites in order to examine a possible predictor function of blackcap song for habitat quality. Several habitat features are responsible for variation in nesting success. These features also correlate with song rate of unmated males. The study indicates a potential role of song rate in the advertisement of territory quality. Furthermore, the data suggest that females use song rates rather than territory quality in mating decisions. The information females may gain about male quality in relation to territory quality are also discussed.     

Modelling the fate of marine turtle hatchlings

The SLIM oceanographic model was used to examine the fate of hatchling flatback turtles (Natator depressus) in the first two weeks of their dispersal starting at Wild Duck Island, a major turtle-nesting site in the central Great Barrier Reef region of Australia. We ran simulations to investigate the effects of spring versus neap tides, hatchling's swimming behaviour during their first three days at sea, and the location of nesting beaches. The model predicted that up to 50% of the turtle hatchlings entering the sea from the windward, southern beach remained after 14 days in shallow, nearshore waters, irrespective of tides and swimming. These waters are turbid and may be inhospitable to hatchlings. In contrast >80% of hatchlings dispersing from the leeward, northern beach were dispersed to deeper water (10-40 m) in a quasi-stationary dispersion core centred around 10 km north of the nesting beach after 14 days and the offshore spread of the turtle plume was enhanced by the hatchling's seaward swimming during the first three days. This was due to the presence of a coastal boundary layer and a stagnation zone around the northern side of island, but not the southern side. The model confirmed that dispersal from eastern Queensland flatback turtle rookeries is restricted to the lagoons and coastal waters, and that water circulation and hatchling's swimming control dispersion. The model explains why more turtles nest on the northern than the southern side. This study highlights the usefulness of oceanographic models to increase knowledge about a cryptic life stage of marine turtles.     

Strong site fidelity and longer internesting interval for solitary nesting olive ridley sea turtles in Brazil

Olive ridley sea turtles display two different types of nesting behavior: in arribada (synchronous mass nesting) or solitarily. Contrarily to arribadas, little has been published about solitary nesters. This study aimed to expand the knowledge on internesting interval and site fidelity of solitary nesting olive ridleys and to test a possible development of arribada nesting behavior. Data were collected in Sergipe (Brazil) over 125 km of beach from 10°30′S/36o23′W to 11°26′S/37o19′W, between nesting seasons 2004/2005 and 2006/2007. From 962 tagged females, 173 were seen renesting. The average internesting interval found was longer (22.35 ± 7.01 days) than previously described, which might relate to lower water temperatures during the internesting period. Olive ridleys at Sergipe showed high nesting site fidelity, with consecutive nesting events occurring in close proximity, non-randomly and dependently of previous events. Most of the consecutive nests were separated by 4.06–5.59 km. Development of arribada nesting behavior was not confirmed.     

Allopaternal care in the fathead minnow,Pimephales promelas: females prefer males with eggs

Summary Some species of fishes with exclusive male parental care exhibit the phenomenon of allopaternal care; that is, some males acquire and care for other males' eggs. We conducted a series of experiments to investigate the dynamics and evolution of allopaternal care in one such species, the fathead minnow (Pimephales promelas). In choosing a nest site, a newly reproductive male tended to take over the nest site of a parental male by evicting the resident male, rather than occupy a physically identical empty nest site. The new male generally cared for the old male's eggs, and in most cases, daily egg survival improved under the new male's care. When males were given a choice among unguarded nest sites, they preferred to occupy nest sites already containing eggs. When eggs were randomly assigned to nesting males, females preferred to spawn with males who had eggs in their nest sites. Thus, it appears that female preference for males with eggs led to the evolution of allopaternal care in the fathead minnow.     

Effects of Illegal Harvest of Eggs on the Population Decline of Leatherback Turtles in Las Baulas Marine National Park,Costa Rica

Abstract: Within 19 years the nesting population of leatherback turtles (Dermochelys coriacea) at Parque Nacional Marino Las Baulas declined from 1500 turtles nesting per year to about 100. We analyzed the effects of fishery bycatch and illegal harvesting (poaching) of eggs on this population. We modeled the population response to different levels of egg harvest (90, 75, 50, and 25%) and the effect of eradicating poaching at different times during the population decline. We compared effects of 90% poaching with those of 20% adult mortality because both of these processes were present in the population at Las Baulas. There was a stepwise decline in number of nesting turtles at all levels of egg harvest. Extirpation times for different levels of poaching ranged from 45 to 282 years. The nesting population declined more slowly and survived longer with 20% adult mortality (146 years) than it did with 90% poaching (45 years). Time that elapsed until poaching stopped determined the average population size at which the population stabilized, ranging from 90 to 420 nesting turtles. Our model predicted that saving clutches lost naturally would restore the population when adult mortality rates were low and would contribute more to population recovery when there were short remigration intervals between nesting seasons and a large proportion of natural loss of clutches. Because the model indicated that poaching was the most important cause of the leatherback decline at Las Baulas, protecting nests on the beach and protecting the beach from development are critical for survival of this population. Nevertheless, the model predicted that current high mortality rates of adults will prevent population recovery. Therefore, protection of the beach habitat and nests must be continued and fishery bycatch must be reduced to save this population.     

Forest Fragmentation Increases Nest Predation in the Eurasian Treecreeper

Abstract:  We used long-term breeding data to monitor the influences of fragmentation and habitat composition at different spatial scales on the reproductive success of Eurasian Treecreepers ( Certhia familiaris ) breeding in nest boxes. We collected data from the same forest patches (2.7–65.1 ha in size) during seven breeding seasons. Nest predation varied considerably over the years and was the primary cause of nesting failure (mean annual rate of 21.6 ± 12.8%). Nest predation explained most of the variation in fledgling production during the study period. Landscape-level fragmentation (radius of 500 m from territory center) affected nest predation more than did fragmentation on the territory scale (radius of 200 m from territory center). In general, nest loss due to predation in fragmented landscapes (32.4%) was almost threefold that of less fragmented (12.0%) landscapes. Of the habitat variables, predation rate correlated positively with the density of edges between forest and open land and with the proportion of sapling stands on the spatial scale of 500 m around a nest. In the core area of a territory (radius of 30 m from territory center), a high density of trees increased the frequency of nest predation. Further, a high proportion of agricultural land close to a nest site increased nest losses of treecreepers, probably because of a high degree of mustelid predation. Our results showed that the spatial scale on which we examined nest predation is important and that even within moderately fragmented landscapes it is possible to detect fragmentation-related nest predation.     

Chemical aspects of mass spawning in corals. II. (-)-Epi-thunbergol,the sperm attractant in the eggs of the soft coral Lobophytum crassum (Cnidaria: Octocorallia)

Sea turtles migrate between feeding and nesting areas that are often geographically separated by hundreds or thousands of kilometers. Observations of their aggregations at sea and at nesting beaches have led to the hypothesis that sea turtles migrate in socially structured groups. While this migratory strategy is common to many marine vertebrates, socially facilitated behavior is not well documented in testudines. In 1990 and 1991, we attached satellite transmitters to olive ridleys (Lepidochelys olivacea Eschscholtz) found ovipositing together during a mass nesting at Nancite Beach, Costa Rica, to determine whether they migrate independently or in groups after they leave the nesting beach. Results showed that the turtles were not spatially associated during the internesting period, were capable of re-establishing themselves as a group during a subsequent nesting emergence, and were not spatially associated during their postnesting migrations to oceanic feeding areas. We suggested that what appear to be socially structured groups of L. olivacea are in fact individual turtles simultaneously using the same habitat.     

Regional and annual variation in plasma steroids and metabolic indicators in female green turtles, <Emphasis Type="Italic">Chelonia mydas</Emphasis>

Variation in environmental conditions at a foraging area or at a nesting rookery has the potential to impact reproductive output of green sea turtles (Chelonia mydas) by affecting food resources or the nesting substrate. In this paper we test whether turtles‘ physiological characteristics reflect variation in relevant environmental conditions. We did this by profiling metabolic and hormonal markers among (1) non-vitellogenic females from three different foraging areas and (2) nesting females from different rookeries and breeding seasons. Among the non-vitellogenic females, the highest plasma triglyceride concentrations (4.29 mmol/l) and the lowest plasma cholesterol concentrations (1.27 mmol/l) were found in non-vitellogenic females residing in Moreton Bay during the El Niño year of 1997. Furthermore, during 1997, these Moreton Bay females had higher plasma triglyceride and lower cholesterol concentrations than those recorded in non-vitellogenic females foraging at Heron Reef (triglyceride 1.22 mmol/l and cholesterol 4.53 mmol/l) and Shoalwater Bay (triglyceride 1.69 mmol/l and cholesterol 3.50 mmol/l) in the same year. Among nesting turtles, those nesting at Raine Island had low mean plasma triglyceride concentrations during the high density 1996 nesting season. For those nesting at Heron Island, the mean triglyceride concentrations were the lowest during the 1997 nesting season. This is the first time that hormone and metabolic markers have been used in concert to compare the physiological condition of nesting and foraging sea turtles and its relationship with the environment. Collectively, our data indicate that variation in the environmental conditions at both foraging and nesting areas are reflected at a physiological level. Moreover, our study indicates that turtles feeding during El Niño years are able to attain higher levels of body condition, and that physiological data combined with morphometric data is a useful proxy for assessing the condition of turtles in foraging areas.     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.     

Mapping risk for nest predation on a barrier island

Barrier islands and coastal beach systems provide nesting habitat for marine and estuarine turtles. Densely settled coastal areas may subsidize nest predators. Our purpose was to inform conservation by providing a greater understanding of habitat-based risk factors for nest predation, for an estuarine turtle. We expected that habitat conditions at predated nests would differ from random locations at two spatial extents. We developed and validated an island-wide model for the distribution of predated Diamondback terrapin nests using locations of 198 predated nests collected during exhaustive searches at Fisherman Island National Wildlife Refuge, USA. We used aerial photographs to identify all areas of possible nesting habitat and searched each and surrounding environments for nests, collecting location and random-point microhabitat data. We built models for the probability of finding a predated nest using an equal number of random points and validated them with a reserve set (N?=?67). Five variables in 9 a priori models were used and the best selected model (AIC weight 0.98) reflected positive associations with sand patches near marshes and roadways. Model validation had an average capture rate of predated nests of 84.14 % (26.17–97.38 %, Q1 77.53 %, median 88.07 %, Q3 95.08 %). Microhabitat selection results suggest that nests placed at the edges of sand patches adjacent to upland shrub/forest and marsh systems are vulnerable to predation. Forests and marshes provide cover and alternative resources for predators and roadways provide access; a suggestion is to focus nest protection efforts on the edges of dunes, near dense vegetation and roads.     

Predicting the Impact of Sea-Level Rise on Caribbean Sea Turtle Nesting Habitat

Abstract:  The projected rise in sea level is likely to increase the vulnerability of coastal zones in the Caribbean, which are already under pressure from a combination of anthropogenic activities and natural processes. One of the major effects will be a loss of beach habitat, which provides nesting sites for endangered sea turtles. To assess the potential impacts of sea-level rise on sea turtle nesting habitat, we used beach profile measurements of turtle nesting beaches on Bonaire, Netherlands Antilles, to develop elevation models of individual beaches in a geographic information system. These models were then used to quantify areas of beach vulnerable to three different scenarios of a rise in sea level. Physical characteristics of the beaches were also recorded and related to beach vulnerability, flooding, and nesting frequency. Beaches varied in physical characteristics and therefore in their vulnerability to flooding. Up to 32% of the total current beach area could be lost with a 0.5-m rise in sea level, with lower, narrower beaches being the most vulnerable. Vulnerability varied with land use adjacent to the beach. These predictions about loss of nesting habitat have important implications for turtle populations in the region.     

Are anthropogenic factors affecting nesting habitat of sea turtles? The case of Kanzul beach,Riviera Maya-Tulum (Mexico)

Marine coast modification and human pressure affects many species, including sea turtles. In order to study nine anthropogenic impacts that might affect nesting selection of females, incubation and hatching survival of loggerhead (Caretta caretta) and green turtle (Chelonia mydas), building structures were identified along a 5.2 km beach in Kanzul (Mexico). A high number of hotels and houses (88; 818 rooms), with an average density of 16.6 buildings per kilometer were found. These buildings form a barrier which prevents reaching the beach from inland, resulting in habitat fragmentation. Main pressures were detected during nesting selection (14.19% of turtle nesting attempts interrupted), and low impact were found during incubation (0.77%) and hatching (4.7%). There were three impacts defined as high: beach furniture that blocks out the movement of hatchlings or females, direct pressure by tourists, and artificial beachfront lighting that can potentially mislead hatchlings or females. High impacted areas showed lowest values in nesting selection and hatching success. Based on our results, we suggest management strategies to need to be implemented to reduce human pressure and to avoid nesting habitat loss of loggerhead and green turtle in Kanzul, Mexico.