First records of flipper beat frequency during sea turtle diving

Depth and flipper movements were simultaneously measured during 23 dives for a free-swimming green turtle (Chelonia mydas) at Ascension Island. A few characteristic dive profiles that have been widely reported in hard-shelled turtles were recorded. Flipper movements revealed that, on dives to midwater, there was generally active swimming, compared to long periods of inactivity on dives to the seafloor. During all dives, there were clear changes in the flipper beat frequency during the descent. On leaving the surface, flippers beats occurred quickly (typically 30-40 beats min⁻¹) and then as the descent continued the frequency declined (typically to about 10-14 beats min⁻¹). These observations match the general pattern reported for other air-breathing divers for increased effort at the start of the descent to overcome initial positive buoyancy.     

Orientation Cues Used by Hatchling Loggerhead Sea Turtles (Caretta caretta L.) During their Offshore Migration

Sea turtle hatchlings ernerge from underground nests, crawl to the ocean and swim out to sea. The orientation cues used to maintain offshore headings are unknown. Our field experiments suggest loggerhead hatchlings on the Atlantic coast of Florida continue on offshore headings by swimming into oceanic swells and wind-generated waves. Dependence on these cues appears complete when hatchlings are 3.0 km from shore. Because waves and swells usually approach from seaward directions, they are generally reliable guideposts.     

The geomagnetic environment in which sea turtle eggs incubate affects subsequent magnetic navigation behaviour of hatchlings

Loggerhead sea turtle hatchlings (Caretta caretta) use regional magnetic fields as open-ocean navigational markers during trans-oceanic migrations. Little is known, however, about the ontogeny of this behaviour. As a first step towards investigating whether the magnetic environment in which hatchlings develop affects subsequent magnetic orientation behaviour, eggs deposited by nesting female loggerheads were permitted to develop in situ either in the natural ambient magnetic field or in a magnetic field distorted by magnets placed around the nest. In orientation experiments, hatchlings that developed in the normal ambient field oriented approximately south when exposed to a field that exists near the northern coast of Portugal, a direction consistent with their migratory route in the northeastern Atlantic. By contrast, hatchlings that developed in a distorted magnetic field had orientation indistinguishable from random when tested in the same north Portugal field. No differences existed between the two groups in orientation assays involving responses to orbital movements of waves or sea-finding, neither of which involves magnetic field perception. These findings, to our knowledge, demonstrate for the first time that the magnetic environment present during early development can influence the magnetic orientation behaviour of a neonatal migratory animal.     

How do goannas find sea turtle nests?

Squamate reptiles rely heavily on visual and chemical cues to detect their prey, so we expected yellow‐spotted goannas (Varanus panoptes) which are predators of sea turtle nests on mainland beaches in northern Australia would use these cues to find sea turtle nests. Ghost crabs (Ocypode ceratophthalmus and Ocypode cordimanus) are also common on Australian sea turtle nesting beaches and frequently burrow into sea turtle nests. However, the potential for ghost crab burrowing activity at sea turtle nests to signal the location of a nest to goannas has not been investigated. Here, we used camera traps and presence of tracks at nests to record goanna activity around selected nests during the incubation period and 10 days after hatchling turtles emerged from their nests. We also recorded the number of ghost crab burrows around nests to evaluate ghost crab activity. Our results indicated that nest discovery by goannas was independent of nest age, but that the nest visitation rate of goannas and crabs increased significantly after a nest had been opened by a goanna or after hatchlings had emerged from the nest. There was no apparent connection between ghost crab burrows into a nest and the likelihood of that nest being predated by goannas.     

Orientation and behaviour of hatchling green turtles (Chelonia mydas) in the sea

To investigate the initial stage of the ‘lost-year puzzle’ of sea turtle ecology, both hatchlings from the natural nesting groung at Tortuguero, Costa Rica and hatchlings from Tortuguero eggs that had hatched in a beach on Bermuda were tracked individually after their departure from these beaches. Of the Bermuda beach most were tracked by swimmers equipped with face mask and flippers and followed by a boat. Tracking off Tortuguero was done by an observer in a following boat. The data showed that non-random departure courses were maintained even when swimming hatchlings and moved over the horizon from all fixed objects on the shore. Observations were made on swimming and diving behaviour and on predator relationships of travelling turtles. The procedures described are useful research techniques and will be used for more extensive tracking in future seasons.     

Does maternal oviposition site influence offspring dispersal to suitable habitat?

Orientation and dispersal to suitable habitat affects fitness in many animals, but the factors that govern these behaviors are poorly understood. In many turtle species, hatchlings must orient and disperse to suitable aquatic habitat immediately after emergence from subterranean nests. Thus, the location of nest sites relative to aquatic habitats ideally should be associated with the direction of hatchling dispersal. At our study site, painted turtles (Chrysemys picta) nest to the west (on an island) and east (on the mainland) of a wetland, which determines the direction that hatchlings must travel to reach suitable aquatic habitat. To determine if hatchling orientation is intrinsically influenced by the location where their mothers nest, we employed a two-part cross-fostering experiment in the field, whereby half the eggs laid in mainland nests were swapped with half the eggs laid in island nests. Moreover, because C. picta hatchlings overwinter inside their nests, we performed a second cross-fostering experiment to fully decouple the effects of (1) the maternally chosen nest location, (2) the embryonic developmental location, and (3) the overwinter location. We released hatchlings into a circular arena in the field and found that turtles generally dispersed in a westerly direction, regardless of the maternally chosen nest location and independent of the locations of embryonic development and overwintering. Although this westerly direction was towards suitable aquatic habitat, we could not distinguish whether naïve hatchling turtles (i) use environmental cues/stimuli to orient their movement, or (ii) have an intrinsic bias to orient west in the absence of stimuli. Nevertheless, these findings suggest that the orientation behavior of naïve hatchling turtles during terrestrial dispersal is not dependent upon the location of maternally-chosen nest sites.     

Compass Orientation During Dispersal of Freshwater Hatchling Snapping Turtles (Chelydra serpentina) and Blanding's Turtles (Emydoidea blandingii)

Freshwater turtle hatchlings primarily use visual cues for orientation while dispersing from nests; however, hatchlings rapidly develop a relationship between a sun or geomagnetic compass and a dispersal target that allows them to maintain an established direction of movement when target habitats are not visible. We examined dispersal patterns of hatchling snapping turtles (Chelydra serpentina) and Blanding's turtles (Emydoidea blandingii) dispersing in large arenas in a mowed field and in dense corn. The dispersal of three categories of hatchlings were examined: (1) naïve individuals (no previous dispersal experience), (2) arena‐experienced (limited dispersal experience in arenas in natural habitat), and (3) natural‐experienced hatchling Blanding's turtles (captured after extensive experience dispersing W in natural habitats toward wetlands). Experienced hatchlings were assigned to treatments consisting of having a magnet or a non‐magnetic aluminum sham or nothing glued to their anterior carapace before release in the corn arena. Dispersal patterns of naïve hatchlings of both species were strongly directional in the field arena with visible target horizons and primarily random in the corn arena where typical target horizons were blocked. When released in corn, dispersal patterns were similar for arena‐experienced hatchlings with magnets or shams attached and differed from their prior dispersal headings in the field arena as naïve hatchlings. Natural‐experienced hatchling Blanding's turtles with and without magnets were able to accurately maintain their prior headings to the WNW while dispersing in the field or corn arenas (i.e., the presence of a magnet did not disrupt their ability to maintain their prior heading). Based on the assumption that no other type of compass exists in hatchlings, we conclude that they were not using a geomagnetic compass, but by default were using sun compass orientation to maintain dispersal headings in dense corn where no typical target habitats were visible.     

Effect of active head movements about the pitch, roll, and yaw axes on human optokinetic afternystagmus

Effects of active head movements about the pitch, roll, or yaw axes on horizontal optokinetic afternystagmas (OKAN) were examined in 16 subjects to test the hypothesis that otolith organ mediated activity induced by a change in head position can couple to the horizontal velocity storage in humans. Active head movements about the pitch axis, forwards or backwards, produced significant OKAN suppression. Pitch forward head movements exerted the strongest effect. Active head movements about the roll axis towards the right also produced OKAN suppression but only if the tilted position was sustained. No suppression was observed following sustained yaw. However, an unsustained yaw left movement after rightward drum rotation significantly enhanced OKAN. Sustained head movement trials did not significantly alter subsequent control trials. In contrast, unsustained movements about the pitch axis, which involve more complex interactions, exerted long-term effects on subsequent control trials. We conclude that otolith organ mediated activity arising from pitch or roll head movements couples to the horizontal velocity storage in humans, thereby suppressing ongoing OKAN. Activity arising from the horizontal canals during an unsustained yaw movement (observed mainly with yaw left), following drum rotation in a direction contralateral to the movement, may also couple to the velocity storage, resulting in increased activity instead of suppression.     

Orientation and swimming behavior of hatchling loggerhead turtles Caretta caretta L. during their offshore migration

Loggerhead turtle Caretta caretta L. hatchlings emerging from nests on the eastern coast of Florida swim offshore toward the Florida Current. Part of the trip is accomplished during an initial 20-h period of swimming (the “frenzy”); the remainder may take a day or more of oriented swimming. Swimming speeds are slower than those of green turtles Chelonia mydas L. Hatchlings are well oriented in an offshore direction when released into the ocean during the day or night. Completion of a crawl down the beach or a plunge into the surf are unnecessary for proper orientation. Immediately after entering the ocean at night, responses to light appear essential for oriented swimming. However, present hypotheses based upon phototropotaxis may not account for orientation later in the migration.     

Sea turtle species vary in their susceptibility to tropical cyclones

Severe climatic events affect all species, but there is little quantitative knowledge of how sympatric species react to such situations. We compared the reproductive seasonality of sea turtles that nest sympatrically with their vulnerability to tropical cyclones (in this study, “tropical cyclone” refers to tropical storms and hurricanes), which are increasing in severity due to changes in global climate. Storm surges significantly decreased reproductive output by lowering the number of nests that hatched and the number of hatchlings that emerged from nests, but the severity of this effect varied by species. Leatherback turtles (Dermochelys coriacea) began nesting earliest and most offspring hatched before the tropical cyclone season arrived, resulting in little negative effect. Loggerhead turtles (Caretta caretta) nested intermediately, and only nests laid late in the season were inundated with seawater during storm surges. Green turtles (Chelonia mydas) nested last, and their entire nesting season occurred during the tropical cyclone season; this resulted in a majority (79%) of green turtle nests incubating in September, when tropical cyclones are most likely to occur. Since this timing overlaps considerably with the tropical cyclone season, the developing eggs and nests are extremely vulnerable to storm surges. Increases in the severity of tropical cyclones may cause green turtle nesting success to worsen in the future. However, published literature suggests that loggerhead turtles are nesting earlier in the season and shortening their nesting seasons in response to increasing sea surface temperatures caused by global climate change. This may cause loggerhead reproductive success to improve in the future because more nests will hatch before the onset of tropical cyclones. Our data clearly indicate that sympatric species using the same resources are affected differently by tropical cyclones due to slight variations in the seasonal timing of nesting, a key life history process.     

Predaceous ants, beach replenishment, and nest placement by sea turtles

Ants known for attacking and killing hatchling birds and reptiles include the red imported fire ant (Solenopsis invicta Buren), tropical fire ant [Solenopsis geminata (Fabr.)], and little fire ant [Wasmannia auropunctata (Roger)]. We tested whether sea turtle nest placement influenced exposure to predaceous ants. In 2000 and 2001, we surveyed ants along a Florida beach where green turtles (Chelonia mydas L.), leatherbacks (Dermochelys coriacea Vandelli), and loggerheads (Caretta caretta L.) nest. Part of the beach was artificially replenished between our two surveys. As a result, mean beach width experienced by nesting turtles differed greatly between the two nesting seasons. We surveyed 1,548 sea turtle nests (2000: 909 nests; 2001: 639 nests) and found 22 ant species. S. invicta was by far the most common species (on 431 nests); S. geminata and W. auropunctata were uncommon (on 3 and 16 nests, respectively). In 2000, 62.5% of nests had ants present (35.9% with S. invicta), but in 2001, only 30.5% of the nests had ants present (16.4% with S. invicta). Turtle nests closer to dune vegetation had significantly greater exposure to ants. Differences in ant presence on turtle nests between years and among turtle species were closely related to differences in nest placement relative to dune vegetation. Beach replenishment significantly lowered exposure of nests to ants because on the wider beaches turtles nested farther from the dune vegetation. Selective pressures on nesting sea turtles are altered both by the presence of predaceous ants and the practice of beach replenishment.     

Invertebrate infestation on eggs and hatchlings of the loggerhead turtle, Caretta caretta, in Dalaman, Turkey

The damage caused by some invertebrates to the eggs and hatchlings of loggerhead turtles, Caretta caretta, was investigated during the summer of 2002 on Dalaman beach, Turkey. The specimens, identified to family or genus levels, from nine families representing seven orders were recorded as infesting nests of loggerhead turtles. The heaviest impacts on loggerhead turtle nests was made by Pimelia sp. (Tenebrionidae, Coleoptera). Twenty-four (36.3%) out of 66 intact loggerhead hatched nests were affected by these larvae. Larval damage by Pimelia sp. was recorded in 188 (10.6%) out of 1773 eggs, but only in two (0.28%) hatchlings. The results show that fewer insects were in the nest the further from vegetation and therefore the relocation of nests from the water's edge to further inland close to vegetation may increase the infestation rate of the eggs.     

The organization of exteroceptive sensory inputs to interneurons of the flight neuropil in locusts

1. Thirty-seven pairs of mesothoracic interneurons respond selectively to visual or ocellar stimuli corresponding to deviations from course in flight, expressed as angular rotation around the three spatial axes. 2. Sensitivities to roll and yaw are very strongly associated. All interneurons showing a directional preference for yaw rotations showed the same preference for roll rotations. A few roll-sensitive cells were not directionally sensitive to yaw. Some interneurons respond exclusively to pitch rotations, most to both pitch and roll/yaw. 3. Approximately equal numbers of interneurons prefer pitch up, pitch down, roll/yaw to the ipsilateral side and roll/yaw to the contralateral side. All four possible combinations of pitch (up or down) with roll/yaw (ipsilateral or contralateral) preferences occur with equal probability. 4. No relationship between neuronal structure and directional properties could be discerned. 5. The average latency of the ocellar EPSPs recorded in the interneurons is not significantly different from the average latency of the ocellar spike in the descending neurons (at the same temperature and in the same ganglion). The average ocellar IPSP latency is 8.5 ms longer. The data support the hypothesis that most EPSPs are derived from monosynaptic inputs from the DNs, and most IPSPs from polysynaptic inputs. A few EPSPs are also derived from polysynaptic inputs. 6. Most of these neurons are sensitive to wind, at least some directionally so, in a manner functionally compatible with their visual or ocellar directionality, and most are excited. Two neurons respond to movement of small objects in the visual field, and 5 to high frequency sound.     

Respiration in neonate sea turtles

The pattern and control of respiration is virtually unknown in hatchling sea turtles. Using incubator-raised turtles, we measured oxygen consumption, frequency, tidal volume, and minute volume for leatherback (Dermochelys coriacea) and olive ridley (Lepidochelys olivacea) turtle hatchlings for the first six days after pipping. In addition, we tested the hatchlings' response to hypercapnic, hyperoxic, and hypoxic challenges over this time period. Hatchling sea turtles generally showed resting ventilation characteristics that are similar to those of adults: a single breath followed by a long respiratory pause, slow frequency, and high metabolic rate. With hypercapnic challenge, both species responded primarily by elevating respiratory frequency via a decrease in the non-ventilatory period. Leatherback resting tidal volume increased with age but otherwise, neither species' resting respiratory pattern nor response to gas challenge changed significantly over the first few days after hatching. At the time of nest emergence, sea turtles have achieved a respiratory pattern that is similar to that of actively diving adults.     

Regional blood flow in sea turtles: implications for heat exchange in an aquatic ectotherm

Despite substantial knowledge on thermoregulation in reptiles, the mechanisms involved in heat exchange of sea turtles have not been investigated in detail. We studied blood flow in the front flippers of two green turtles, Chelonia mydas, and four loggerhead turtles, Caretta caretta, using Doppler ultrasound to assess the importance of regional blood flow in temperature regulation. Mean blood flow velocity and heart rate were determined for the water temperature at which the turtles were acclimated (19.3 degrees-22.5 degrees C) and for several experimental water temperatures (17 degrees-32 degrees C) to which the turtles were exposed for a short time. Flipper circulation increased with increasing water temperature, whereas during cooling, flipper circulation was greatly reduced. Heart rate was also positively correlated with water temperature; however, there were large variations between individual heart rate responses. Body temperatures, which were additionally determined for the two green turtles and six loggerhead turtles, increased faster during heating than during cooling. Heating rates were positively correlated with the difference between acclimation and experimental temperature and negatively correlated with body mass. Our data suggest that by varying circulation of the front flippers, turtles are capable of either transporting heat quickly into the body or retaining heat inside the body, depending on the prevailing thermal demands.     

A novel hypothesis for the adaptive maintenance of environmental sex determination in a turtle

Temperature-dependent sex determination (TSD) is widespread in reptiles, yet its adaptive significance and mechanisms for its maintenance remain obscure and controversial. Comparative analyses identify an ancient origin of TSD in turtles, crocodiles and tuatara, suggesting that this trait should be advantageous in order to persist. Based on this assumption, researchers primarily, and with minimal success, have employed a model to examine sex-specific variation in hatchling phenotypes and fitness generated by different incubation conditions. The unwavering focus on different incubation conditions may be misplaced at least in the many turtle species in which hatchlings overwinter in the natal nest. If overwintering temperatures differentially affect fitness of male and female hatchlings, TSD might be maintained adaptively by enabling embryos to develop as the sex best suited to those overwintering conditions. We test this novel hypothesis using the painted turtle (Chrysemys picta), a species with TSD in which eggs hatch in late summer and hatchlings remain within nests until the following spring. We used a split-clutch design to expose field-incubated hatchlings to warm and cool overwintering (autumn–winter–spring) regimes in the laboratory and measured metabolic rates, energy use, body size and mortality of male and female hatchlings. While overall mortality rates were low, males exposed to warmer overwintering regimes had significantly higher metabolic rates and used more residual yolk than females, whereas the reverse occurred in the cool temperature regime. Hatchlings from mixed-sex nests exhibited similar sex-specific trends and, crucially, they were less energy efficient and grew less than same-sex hatchlings that originated from single-sex clutches. Such sex- and incubation-specific physiological adaptation to winter temperatures may enhance fitness and even extend the northern range of many species that overwinter terrestrially.     

Acquisition of Salmonella flora by turtle hatchlings on commercial turtle farms

A commercial turtle pond in South Louisiana was studied to identify the mechanism by which turtle hatchlings acquire Salmonella flora. The visceral organs and mature eggs removed from 31 adult gravid female turtles over the course of two egg-laying seasons and from 37 adult females during one winter dormant period were examined bacteriologically for Salmonella. Pond water, egg nest soil, and hatchlings produced by eggs removed from the oviducts and nest soil were also tested. Eighty-eight turtles hatched from eggs removed from the oviducts of 15 turtles at necropsy did not excrete or harbor systemically Salmonella, nor were these pathogens isolated from ovarian tissue or immature eggs. The findings suggest transovarian transmission of these pathogens does not occur frequently. Turtles hatched from eggs retrieved from soil nests 1 to 2 h after deposition harbor and excrete these organisms. This result coupled with the isolation of these pathogens from the cloaca, colon contents, and bursal fluid from 18 females captured in the act of egg laying supports the cloaca to egg and nest soil to egg mode for salmonellae infection in the resultant hatchling. Salmonella arizonae and Salmonella serogroups B, C2, and E1 were isolated from the cloaca, colon contents, pond water, and nest soil, and were excreted by hatchlings produced from eggs removed from the soil nests. These same serogroups were isolated from the colon contents of 19 of 37 females tested during the dormant period, suggesting the salmonellae persist in the pond environment in the adult throughout the year.     

Metabolic heat estimation of the sea turtle Lepidochelys olivacea embryos

Several studies have reported the importance of metabolic heat on the increment of temperature in the sea turtle nests; however, the metabolic heat has not been calculated for sea turtle eggs. In this study, the metabolic heat generated by embryos of the sea turtle Lepidochelys olivacea was estimated from a thermal balance model by means of three measured temperatures—one in the center of the nest, and the others in the sand above and beside the nest. An experiment was conducted with a sample of 100 eggs from a Lepidochelys olivacea nest collected in the Baja Peninsula, Mexico. The results showed that during the incubation period, no metabolic heat was detected before day 19 but it increased from that day until a maximum of 0.84 W at day 34, when the incubation process was interrupted due to rain. This value corresponds to 31 emerged hatchlings. The novel model is a suitable framework to predict the temperature and metabolic heat within the nest.     

Fuel use and corticosterone dynamics in hatchling green sea turtles (Chelonia mydas) during natal dispersal

During their natal dispersal hatchling sea turtles depart their nest, beach and inshore areas quickly to move into offshore developmental habitat using their finite energy stores. Patterns of fuel use and endocrine responses that could facilitate hatchling sea turtle dispersal activity are poorly understood. This study, examined aspects of intermediary metabolism by measuring plasma fuel use and an endocrine response of hatchling green turtles (Chelonia mydas) during terrestrial and aquatic activity coinciding with natal dispersal. Specifically, we measured plasma concentrations of glucose, non-esterised free fatty acids and protein to gauge the contributions of carbohydrate, lipid and protein metabolism for fuelling natal dispersal. In addition, we measured plasma levels of the steroid hormone corticosterone (CORT) a hormone implicated in regulating a number of metabolic events associated with migration and energy use in vertebrates. During terrestrial activity, hatchlings ascended through the sand from their nests and exhibited significant increases in plasma CORT and lactate indicating intense periods of anaerobic activity. During swimming, all plasma metabolites, with the exception of plasma protein, peaked between 1 and 4 h post-beginning swimming activity. Plasma CORT peaked at between 3 and 5 h of swimming activity. These plasma concentrations are consistent with intensive activity inducing catabolism of carbohydrate, lipid and protein stores to support prolonged activity. These results are similar to other vertebrates and suggest a relatively uniform cascade of physiological processes during such arduous migratory events.