Polyandry in a marine turtle : females make the best of a bad job

The female perspective on reproductive strategies remains one of the most active areas of debate in biology. Even though a single mating is often sufficient to satisfy the fertilization needs of most females and the act of further mating incurs costs, multiple paternity within broods or clutches is a common observation in nature. Direct or indirect advantage to females is the most popular explanation. However, the ubiquity of this explanation is being challenged by an increasing number of cases for which benefits are not evident. For the first time, we test possible fitness correlates of multiple paternity in a marine turtle, an organism that has long attracted attention in this area of research. Contrary to the wide-spread assumption that multiple mating by female marine turtles confers fitness benefits, none were apparent. In this study, the environment played a far stronger role in determining the success of clutches than whether paternity had been single or multiple. A more likely explanation for observations of multiply sired clutches in marine turtles is that these are successful outcomes of male coercion, where females have conceded to superfluous matings as a compromise. Thus, multiple matings by female marine turtles may be a form of damage control as females attempt to make the best of a bad job in response to male harassment.

The implications of adult morphology for clutch size in the flatback turtle (Natator depressa)

When the mean adult length and mean clutch volume of marine turtles are examined, a clear pattern for larger species to lay larger clutches is evident, in accord with predictions that female size constrains the available space for carrying eggs. However, when compared with this general trend, the volume of clutches laid by flatback turtles (Natator depressa) are smaller than expected. The implication is that the unusually flat morphology of flatback turtles, provides an additional constraint on their egg carrying capacity.

The importance of sand albedo for the thermal conditions on sea turtle nesting beaches

At Ascension Island and Cyprus, major nesting areas for green turtles (Chelonia mydas) in the Atlantic and Mediterranean, respectively, visual inspection shows some beaches are light in colour while others are darker. We objectively measured the albedo of the sand on different beaches, i.e. the percentage of the incident solar radiation that was reflected from the sand surface. At sites where albedo was recorded, we also measured the temperature of the sand at nest depths. At both rookeries, the sand temperature was markedly higher on darker beaches due to greater absorption of the incident solar radiation over the diurnal cycle. Temperature loggers buried at nest depths revealed seasonal changes in temperature on both islands, but showed that the lowest temperatures found on the darker beaches rarely dropped below the highest temperatures on the lighter beaches. Sea turtles exhibit temperature-dependent sex determination. Since sand albedo is a major avenue for the production of a range of incubation temperatures on both islands, it will also have profound implications for hatchling sex ratios. In comparison with both Ascension Island and Cyprus, for samples collected from sea turtle rookeries around the world there was an even greater range in sand albedo values. This suggests that sand albedo, a factor that has previously received little consideration, will have profound implications for nest temperatures, and hence hatchling sex ratios, for other populations and species.

Movement patterns for a critically endangered species, the Leatherback Turtle (Dermochelys coriacea), linked to foraging success and population status

Foraging success for pelagic vertebrates may be revealed by horizontal and vertical movement patterns. We show markedly different patterns for leatherback turtles in the North Atlantic versus Eastern Pacific, which feed on gelatinous zooplankton that are only occasionally found in high densities. In the Atlantic, travel speed was characterized by two modes, indicative of high foraging success at low speeds (<15 km d−1) and transit at high speeds (20–45 km d−1). Only a single mode was evident in the Pacific, which occurred at speeds of 21 km d−1 indicative of transit. The mean dive depth was more variable in relation to latitude but closer to the mean annual depth of the thermocline and nutricline for North Atlantic than Eastern Pacific turtles. The most parsimonious explanation for these findings is that Eastern Pacific turtles rarely achieve high foraging success. This is the first support for foraging behaviour differences between populations of this critically endangered species and suggests that longer periods searching for prey may be hindering population recovery in the Pacific while aiding population maintenance in the Atlantic.

The implications of lung-regulated buoyancy control for dive depth and duration

Among air-breathing divers, control of buoyancy through lung volume regulation may be most highly developed in marine turtles. In short, the turtle lung may serve a dual role as both an oxygen store and in buoyancy control. A simple model is developed to show that, for turtles diving up to the maximum depth at which they can still use their lungs to attain neutral buoyancy, the total oxygen store will increase greatly with dive depth, and hence a corresponding increase in dive duration is predicted. Time–depth recorders attached to free-living green turtles (Chelonia mydas) at Ascension Island confirmed a marked increase in dive duration with depth, with the gradient of this relationship being >10 times that seen in diving birds and mammals. Consistent with the prediction that the lungs serve a dual role, we found that, when lead weights were added to some turtles to increase their specific gravity, the mean depth of dives decreased, but for dives to the same depth, weighted animals dived for longer. The depth distribution of green turtles seems to be generally constrained by the maximum depth at which they can still attain close to neutral buoyancy.

Behavioural plasticity in a large marine herbivore: contrasting patterns of depth utilisation between two green turtle (Chelonia mydas) populations

We used time-depth recorders to measure depth utilisation in gravid green turtles (Chelonia mydas) during the internesting period at northern Cyprus (Mediterranean), a nesting area where individuals feed, and at Ascension Island (mid-Atlantic), a nesting area where individuals fast. There were contrasting patterns of depth utilisation between the two sites, illustrating that the behaviour of this species is shaped by local conditions. For example, the amount of time spent shallower than 4 m was 90% at Cyprus but only 31% at Ascension Island, and there was a clear difference between the mean depth at Cyprus (2.7 m, n=9 internesting intervals) versus Ascension Island (9.5 m, n=6 internesting intervals) (t 5=5.92, P=0.002). At Cyprus, turtles spent the greatest percentage of their time at very shallow depths, where surveys reveated a high abundance of seagrass on which this population feeds. In contrast, the deeper distribution at Ascension Island may reflect the preferred depth for resting on the seabed.

Oceanic long-distance navigation : do experienced migrants use the earth's magnetic field?

Albatrosses and sea turtles are known to perform extremely long-distance journeys between disparate feeding areas and breeding sites located on small, isolated, oceanic islands or at specific coastal sites. These oceanic journeys, performed mainly over or through apparently featureless mediums, indicate impressive navigational abilities, and the sensory mechanisms used are still largely unknown. This research used three different approaches to investigate whether bi-coordinate navigation based on magnetic field gradients is likely to explain the navigational performance of wandering albatrosses in the South Atlantic and Indian Oceans and of green turtles breeding on Ascension Island in the South Atlantic Ocean. The possibility that magnetic field parameters can potentially be used in a bi-coordinate magnetic map by wandering albatrosses in their foraging area was investigated by analysing satellite telemetry data published in the literature. The possibilities for using bi-coordinate magnetic navigation varied widely between different areas of the Southern Oceans, indicating that a common mechanism, based on a bi-coordinate geomagnetic map alone, was unlikely for navigation in these areas. In the second approach, satellite telemetry was used to investigate whether Ascension Island green turtles use magnetic information for navigation during migration from their breeding island to foraging areas in Brazilian coastal waters. Disturbing magnets were applied to the heads and carapaces of the turtles, but these appeared to have little effect on their ability to navigate. The only possible effect observed was that some of the turtles with magnets attached were heading for foraging areas slightly south of the control turtles along the Brazilian coast. In the third approach, breeding female green turtles were deliberately displaced in the waters around Ascension Island to investigate which cues these turtles might use to locate and return to the island; the results suggested that cues transported by wind might be involved in the final stages of navigation.

Asynchronous emergence by loggerhead turtle (Caretta caretta) hatchlings

For many decades it has been accepted that marine turtle hatchlings from the same nest generally emerge from the sand together. However, for loggerhead turtles (Caretta caretta) nesting on the Greek Island of Kefalonia, a more asynchronous pattern of emergence has been documented. By placing temperature loggers at the top and bottom of nests laid on Kefalonia during 1998, we examined whether this asynchronous emergence was related to the thermal conditions within nests. Pronounced thermal variation existed not only between, but also within, individual nests. These within-nest temperature differences were related to the patterns of hatchling emergence, with hatchlings from nests displaying large thermal ranges emerging over a longer time-scale than those characterised by more uniform temperatures.

Thermal conductivity of sand and its effect on the temperature of Loggerhead Sea Turtle (Caretta Caretta) nests

The conductivity of sand at a depth of 30–50 cm was measured at 15 sites on the beach at Captiva Island in south-west Florida which is used by nesting loggerhead turtles (Caretta caretta). The mean daily temperature of the sand was correlated with conductivity at the same depth measured the same day (r=0·611). When day to day variation was removed the correlation between nest temperature and conductivity increased to 0·694. The sand was highly variable in its grain structure. The dominant variability (80·6%) was redescribed by the first two principal components of a Principal Components Analysis (PCA). These two components were influenced mostly by percentages of large (> 1 mm) and small (< 500 μm) grains respectively. Conductivity was strongly correlated with the grain structure of the sand. The first three principal components describing sand grain structure, explained 84·1% of the variation in conductivity. Moisture content of the sand (always < 5%) was not an important factor. Sites dominated by larger grains generally had poorer conductivity and were cooler. Comparisons of eight nests to seven adjacent random sites revealed no strong evidence for directional selection in nest placement relative to sand conductivity. The variance in conductivities recorded at nests was also not significantly different from the variance at random sites.

Inter- and intra-beach thermal variation for Green Turtle nests on Ascension Island, South Atlantic

Nest temperatures for green turtles (Chelonia mydas) nesting on Ascension Island, South Atlantic (7°57'S 14°22'W), were examined. Temperature probes were placed into nests on two beaches, Long Beach (26 nests) and North East Bay (8 nests). Within these beaches there was relatively little thermal variation (SD of nest temperature was 0.32°C for Long Beach and 0.30°C for North East Bay). To examine inter-beach thermal variation temperature probes were buried at 55 cm on 12 beaches. Inter-beach thermal variation was large and was related to the beach albedo with the darkest beach (albedo, 016) being 4.2°C warmer than the lightest coloured beach (albedo, 0.73).

Behaviour and buoyancy regulation in the deepest-diving reptile : the leatherback turtle

In the face of the physical and physiological challenges of performing breath-hold deep dives, marine vertebrates have evolved different strategies. Although behavioural strategies in marine mammals and seabirds have been investigated in detail, little is known about the deepest-diving reptile – the leatherback turtle (Dermochelys coriacea). Here, we deployed tri-axial accelerometers on female leatherbacks nesting on St Croix, US Virgin Islands, to explore their diving strategy. Our results show a consistent behavioural pattern within dives among individuals, with an initial period of active swimming at relatively steep descent angles (∼–40 deg), with a stroke frequency of 0.32 Hz, followed by a gliding phase. The depth at which the gliding phase began increased with the maximum depth of the dives. In addition, descent body angles and vertical velocities were higher during deeper dives. Leatherbacks might thus regulate their inspired air-volume according to the intended dive depth, similar to hard-shelled turtles and penguins. During the ascent, turtles actively swam with a stroke frequency of 0.30 Hz but with a low vertical velocity (∼0.40 ms–1) and a low pitch angle (∼+26 deg). Turtles might avoid succumbing to decompression sickness (‘the bends’) by ascending slowly to the surface. In addition, we suggest that the low body temperature of this marine ectotherm compared with that of endotherms might help reduce the risk of bubble formation by increasing the solubility of nitrogen in the blood. This physiological advantage, coupled with several behavioural and physical adaptations, might explain the particular ecological niche the leatherback turtle occupies among marine reptiles.

Conservation hotspots : implications of intense spatial area use by breeding male and female loggerheads at the Mediterranean’s largest rookery

The implementation of appropriate protection measures for endangered species in protected areas requires knowledge of their fine-scale habitat use. In May and June of 2006 and 2007, we used GPS loggers (some linked to the Argos system) and a conventional Argos transmitter to track male and female loggerhead turtles Caretta caretta in the vicinity of the breeding area of Laganas Bay within the National Marine Park of Zakynthos, Greece. We obtained (1) 9681 useable locations (mean: 1383 locations ind.–1; range: 519 to 2198 locations) from Tracktag GPS loggers attached to 7 females for a mean duration of 34 d (range: 17 to 52 d); (2) 1245 useable locations (mean: 311 locations ind.–1; range: 38 to 1110 locations) from 4 males fitted with Fastloc Argos tags for a mean duration of 29 d (range: 3 to 51 d) and (3) 100 locations from 1 male fitted with a conventional Argos satellite tag tracked for 128 d. GPS data indicated that before the onset of nesting, both males and females primarily used an area within 500 m of the shore along a core 9 km stretch of coastline, where existing protective legislation requires strengthening. Our observations suggest that a 76.7% female-biased operational sex ratio, measured previously from in-water surveys, may represent a realistic sex ratio estimate in the period before nesting starts. In the first month following the onset of nesting, female spatial distribution remained similar, whereas most males departed for distant areas presumably to forage. Our study provides quantitative evidence of the need to improve the management planning and conservation measures to protect sea turtles in a coastal breeding area, and new insights on male turtle migration.

Indicators of the impact of climate change on migratory species

The Bonn Convention on the Conservation of Migratory Species of Wild Animals adopted a Resolution in 2005 recognising the impacts of climate change on migratory species. It called on Contracting Parties to undertake more research to improve our understanding of these impacts and to implement adaptation measures to reduce foreseeable adverse effects. Given the large diversity of taxa and species affected by climate change, it is impossible to monitor all species and effects thereof. However, it is likely that many of the key ecological and physical processes through which climate change may impact wildlife could be monitored using a suite of indicators, each comprising parameters of species/populations or groups of species as proxies for wider assemblages, habitats and ecosystems. Herein, we identify a suite of 17 indicators whose attributes could reveal negative impacts of climate change on the global status of migratory species: 4 for birds, 4 for marine mammals, 2 for sea turtles, 1 for fish, 3 for land mammals and 3 for bats. A few of these indicators would be relatively straightforward to develop, but most would require additional data collation, and in many cases methodological development. Choosing and developing indicators of the impacts of climate change on migratory species is a challenge, particularly with endangered species, which are subject to many other pressures. To identify and implement conservation measures for these species, indicators must account for the full ensemble of pressures, and link to a system of alerts and triggers for action.

Travelling through a warming world: climate change and migratory species

Long-distance migrations are among the wonders of the natural world, but this multi-taxon review shows that the characteristics of species that undertake such movements appear to make them particularly vulnerable to detrimental impacts of climate change. Migrants are key components of biological systems in high latitude regions, where the speed and magnitude of climate change impacts are greatest. They also rely on highly productive seasonal habitats, including wetlands and ocean upwellings that, with climate change, may become less food-rich and predictable in space and time. While migrants are adapted to adjust their behaviour with annual changes in the weather, the decoupling of climatic variables between geographically separate breeding and non-breeding grounds is beginning to result in mistimed migration. Furthermore, human land-use and activity patterns will constrain the ability of many species to modify their migratory routes and may increase the stress induced by climate change. Adapting conservation strategies for migrants in the light of climate change will require substantial shifts in site designation policies, flexibility of management strategies and the integration of forward planning for both people and wildlife. While adaptation to changes may be feasible for some terrestrial systems, wildlife in the marine ecosystem may be more dependent on the degree of climate change mitigation that is achievable.

Acceleration data reveal the energy management strategy of a marine ectotherm during reproduction

1. Maintaining a high and stable body temperature is often critical for female ectotherms during reproduction. Yet this strategy may be energetically costly, and therefore challenging, during this period of already high-energy demand. 2. Here, the 6-week deployment of tri-axial accelerometers (n = 6) on a marine ectotherm, the loggerhead turtle (Caretta caretta), reproducing at the northern limit of the species’ breeding range (i.e. in a thermally dynamic environment) revealed the behavioural mechanisms underlying its energy management strategy during the breeding season. 3. The estimated activity levels of female loggerheads using overall dynamic body acceleration (ODBA) were high during the breeding season, suggesting that marine turtles may not be able to remain inactive for long periods in the same manner as terrestrial ectotherms, because of the thermally dynamic nature of their environment. 4. However, activity levels were not constant throughout the season, being impacted by both ambient water temperature and female reproductive status. In cold water at the beginning of the nesting season, high levels of activity suggested that females behaviourally thermoregulated by seeking out warm water patches along the shoreline. Interactions with male turtles (courtship and/or avoidance) may also explain this high level of activity. As sea temperatures warmed up and the amount of energy devoted to reproduction probably increased, the turtles spent more time resting during long sequential flat-bottomed dives, and reduced any unnecessary locomotory activity. 5. Turtles may therefore adjust their activity patterns in response to seasonal variations in abiotic (i.e. ambient temperature) and biotic (i.e. reproductive status) factors. This may help minimize activity-linked metabolic rate and maximize reproductive output over a season while breeding in thermally dynamic environments. 6. A mechanistic model gave support to these empirical results. The model revealed that actively maintaining high and stable body temperature is of clear benefit to female turtles at temperate breeding sites. While energetically costly, such active thermoregulatory behaviour may speed up egg maturation, allowing turtles to initiate nesting earlier in the season, and hence maximize reproductive output.