Effects of temperature and moisture during incubation on carcass composition of hatchling snapping turtles (Chelydra serpentina)

Summary Flexible-shelled eggs of common snapping turtles (Chelydra serpentina) were incubated on each of two substrates (vermiculite, sand) at each of three temperatures (26.0°C, 28.5°C, 31.0°C) and three moisture regimes (wet, intermediate, dry). Embryos developing in cool, wet environments mobilized the largest amounts of protein from their yolk and attained the largest size before hatching, whereas turtles developing in warm, dry environments mobilized the smallest quantities of protein and were the smallest in body size at hatching. Embryos on wet substrates mobilized more lipid from their yolk than did embryos on dry media, but ambient temperature had no demonstrable influence on patterns of lipid mobilization. The total reserve of neutral lipid available in residual yolk plus carcass to sustain neonates in the interval prior to the beginning of feeding was largest in hatchlings from dry environments and smallest in animals from wet environments, but was unaffected by temperature during incubation. Hydration of tissues in hatchlings was higher when incubation was in cool, moist conditions than when incubation was in warm, dry settings, thereby indicating that some of the effects of moisture and temperature on mobilization of nutrients by embryos may be mediated by differences in intracellular water. Patterns of response to temperature and moisture recorded for turtles emerging from eggs on sand were similar to those recorded for hatchlings on vermiculite, so no important conclusion would have been affected by incubating eggs on one medium instead of the other.     

Temperature during embryonic and juvenile development influences growth in hatchling snapping turtles,Chelydra serpentina

Embryonic temperature influenced subsequent growth in juvenile snapping turtles, Chelydra serpentina: incubation temperatures of 24 and 26.5°C enhanced growth relative to a temperature of 29°C. Although embryonic temperature normally determines gonadal sex in this species, experimental manipulations revealed that temperature effects on growth were independent of sex. Ambient temperature also affected growth: juvenile turtles grew slowly in a cool (19°C) versus a warm (28°C) environment. In a parallel experiment, turtles from different embryonic temperatures displayed different patterns of temperature choice in response to nutritional status or time of day. We tentatively conclude that embryonic temperature has both direct and indirect (i.e., through temperature choice) effects on growth in snapping turtles.     

Behavioral responses of hatchling painted turtles (Chrysemys picta) and snapping turtles (Chelydra serpentina) at subzero temperatures

We monitored behavioral responses of cold-acclimated hatchling painted turtles (Chrysemys picta) indigenous to Nebraska and hatchling snapping turtles (Chelydra serpentina) indigenous to Nebraska and Arkansas during cooling (0.1°C/min) to temperatures as low as −19°C. All turtles made exploratory movements during cooling and locomotion occurred at temperatures as low as −2 to −4°C, but C. picta maintained relatively higher levels of locomotor activity than C. serpentina, and no differences in motility occurred between northern and southern groups of C. serpentina. Slow movements of the head and limbs were observed in supercooled hatchling C. picta at temperatures as low as −10°C, whereas at about −5°C, C. serpentina exhibited an increase in spontaneous motor activity followed by muscle contracture, immobility, and spontaneous freezing. C. picta spontaneously froze at about −16°C without exhibiting cold contracture, suggesting that they are better adapted to survive exposure to extreme cold.     

Swimming performance of hatchling green turtles is affected by incubation temperature

In an experiment repeated for two separate years, incubation temperature was found to affect the body size and swimming performance of hatchling green turtles (Chelonia mydas). In the first year, hatchlings from eggs incubated at 26°C were larger in size than hatchlings from 28 and 30°C, whilst in the second year hatchlings from 25.5°C were similar in size to hatchings from 30°C. Clutch of origin influenced the size of hatchlings at all incubation temperatures even when differences in egg size were taken into account. In laboratory measurements of swimming performance, in seawater at 28°C, hatchlings from eggs incubated at 25.5 and 26°C had a lower stroke rate frequency and lower force output than hatchlings from 28 and 30°C. These differences appeared to be caused by the muscles of hatchlings from cooler temperatures fatiguing at a faster rate. Clutch of origin did not influence swimming performance. This finding that hatchling males incubated at lower temperature had reduced swimming ability may affect their survival whilst running the gauntlet of predators in shallow near-shore waters, prior to reaching the relative safety of the open sea.     

Physiological ecology of overwintering in hatchling turtles

Temperate species of turtles hatch from eggs in late summer. The hatchlings of some species leave their natal nest to hibernate elsewhere on land or under water, whereas others usually remain inside the nest until spring; thus, post-hatching behavior strongly influences the hibernation ecology and physiology of this age class. Little is known about the habitats of and environmental conditions affecting aquatic hibernators, although laboratory studies suggest that chronically hypoxic sites are inhospitable to hatchlings. Field biologists have long been intrigued by the environmental conditions survived by hatchlings using terrestrial hibernacula, especially nests that ultimately serve as winter refugia. Hatchlings are unable to feed, although as metabolism is greatly reduced in hibernation, they are not at risk of starvation. Dehydration and injury from cold are more formidable challenges. Differential tolerances to these stressors may explain variation in hatchling overwintering habits among turtle taxa. Much study has been devoted to the cold-hardiness adaptations exhibited by terrestrial hibernators. All tolerate a degree of chilling, but survival of frost exposure depends on either freeze avoidance through supercooling or freeze tolerance. Freeze avoidance is promoted by behavioral, anatomical, and physiological features that minimize risk of inoculation by ice and ice-nucleating agents. Freeze tolerance is promoted by a complex suite of molecular, biochemical, and physiological responses enabling certain organisms to survive the freezing and thawing of extracellular fluids. Some species apparently can switch between freeze avoidance or freeze tolerance, the mode utilized in a particular instance of chilling depending on prevailing physiological and environmental conditions.     

Availability of water affects organ growth in prenatal and neonatal snapping turtles (Chelydra serpentina)

We manipulated the amount of water that was available to prenatal and neonatal snapping turtles (Chelydra serpentina) in order to assess the impact of water on growth by different organs in these animals. Three treatments were used: (1) turtles that completed their incubation on a wet substrate, (2) turtles that completed their incubation on a dry substrate, and (3) turtles that spent a few days in water after completing incubation on a dry substrate. Turtles hatching on a dry substrate (treatment 2) were smaller than animals in the other two treatments (which did not differ in size), so data for mass of different organs were adjusted by ANCOVA to remove effects of body size. Scaled masses of liver, stomach, lungs, kidneys, and small intestine did not differ between turtles emerging in wet environments and those hatching in dry environments, but hearts of turtles hatching in dry settings were substantially larger than those of animals hatching in wet ones. Thus, the mass of most organs in turtles developing in wet and dry environments scaled to body size, whereas the heart was hypertrophied in embryos developing in dry environments. Turtles that spent a few days in water after hatching from eggs in dry environments grew rapidly in size, and the increase in body size was accompanied by disproportionately rapid growth in the liver, stomach, lungs, kidneys, and small intestine. The heart did not increase in size during this period, despite the substantial increase in body mass over that at hatching. The enlarged heart of turtles hatching on dry substrates may have been caused by a circulatory hypovolemia late in incubation; the rapid growth of organs other than the heart when these animals were placed in water may reflect a release from constraints on growth once circulatory volume was restored. Accepted: 2 November 1999     

Locomotion in hatchling leatherback turtles Dermochelys coriacea

Hatchling leatherback turtles can only swim forwards, and employ synchronized beating of the forelimbs whether swimming slowly or quickly. The hind limbs make no contribution to propulsion. Effectively, the hatchlings have two swimming speeds; subsurface and fast (30 cm s^-1) or surfaced and slow (8 cm s^-1). Intermediate velocities are transitory; the hatchlings were never seen to rest without movement, nor did they exhibit gliding of the type seen in green turtles. During fast ('vigorous') swimming, power is developed on both the upstroke and downstroke of the limb cycle. During slow swimming, power is only developed during the upstroke—a consequence of the orientation of the axis of limb beat which is opposite in direction to that of cheloniid sea turtles. Terrestrial locomotion is laboured and features an unstable gait which involves simultaneous movement of all four limbs and forward overbalancing during each limb cycle.     

Thermal control of hatchling emergence patterns in marine turtles

The emergence patterns of green turtle (Chelonia mydas) hatchlings on two beaches on Ascension Island, South Atlantic were monitored and related to thermal patterns in the sand at 10, 20, 30 and 40 cm depth. A total of 6001 hatchlings were recorded emerging on Long Beach, and 3171 emerged on North East Bay during the study period. No significant difference was observed in the temporal pattern of hatchling emergence among nights, or between the two beaches. Hatchling emergence predominantly occurred at night with over 93% of hatchlings emerging during the hours of darkness. Almost all hatchlings emerging in daylight suffer predation by the Ascension frigatebird (Fregata aquila). Counts of frigatebirds both above the study beaches and offshore were highest just after sunrise, with a smaller peak prior to sunset, when frigatebirds were found to predate hatchlings emerging, crawling down the beach or detected in inshore waters. The likely thermal cues controlling hatchling emergence were investigated (temperature at different depths, thermal gradients in the sand and temperature change). The most plausible thermal factor appears to be the change of temperature at superficial sand depths, with hatchling emergence inhibited when subsurface sand temperatures were increasing. This simple mechanism is likely to ensure predominantly nocturnal hatchling emergence regardless of sand albedo, seasonality or latitude as long as night is relatively cooler than day.     

Transmission of Haemogregarina balli from painted turtles to snapping turtles through the leech Placobdella ornata.

Six leeches (Placobdella ornata) were allowed to feed on a painted turtle (Chrysemys picta marginata) infected with Haemogregarina balli and subjected to a period of diapause before being allowed to feed on 2 laboratory-reared snapping turtles (Chelydra serpentina). Weekly examination of thin blood films revealed infections of the turtles at 130 days postfeeding. These observations provide support for broad host specificity of hemogregarine parasites of chelonians.     

Measuring energy expenditure in sub-adult and hatchling sea turtles via accelerometry

Measuring the metabolic of sea turtles is fundamental to understanding their ecology yet the presently available methods are limited. Accelerometry is a relatively new technique for estimating metabolic rate that has shown promise with a number of species but its utility with air-breathing divers is not yet established. The present study undertakes laboratory experiments to investigate whether rate of oxygen uptake (o₂) at the surface in active sub-adult green turtles Chelonia mydas and hatchling loggerhead turtles Caretta caretta correlates with overall dynamic body acceleration (ODBA), a derivative of acceleration used as a proxy for metabolic rate. Six green turtles (25–44 kg) and two loggerhead turtles (20 g) were instrumented with tri-axial acceleration logging devices and placed singly into a respirometry chamber. The green turtles were able to submerge freely within a 1.5 m deep tank and the loggerhead turtles were tethered in water 16 cm deep so that they swam at the surface. A significant prediction equation for mean o₂ over an hour in a green turtle from measures of ODBA and mean flipper length (R² = 0.56) returned a mean estimate error across turtles of 8.0%. The range of temperatures used in the green turtle experiments (22–30°C) had only a small effect on o₂. A o₂-ODBA equation for the loggerhead hatchling data was also significant (R² = 0.67). Together these data indicate the potential of the accelerometry technique for estimating energy expenditure in sea turtles, which may have important applications in sea turtle diving ecology, and also in conservation such as assessing turtle survival times when trapped underwater in fishing nets.     

Cardiovascular correlates of exercise in snapping turtles, Chelydra serpentina

1. Heart rate increased with a rise in body temperature (10-30 degrees C) and with induced physical exercise in snapping turtles. 2. Maximum heart rate increment occurred at 30 degrees C. 3. Standard oxygen pulse did not change with a rise in temperature. 4. Oxygen pulse during exercise and oxygen pulse increment were maximal at 10 degrees C and minimal at 20 degrees C. 5. The increase in heart rate with exercise accounted for only 9-22% of the increase in oxygen transport during activity; the remainder was provided by a rise in cardiac stroke volume and/or A-V difference.     

Effects of acclimation and egg-incubation temperature on selected temperature by hatchling western painted turtles (Chrysemys picta bellii)

The ability of hatchling turtles to detect environmental temperature differences and to effectively select preferred temperature is a function that critically impacts survival. In some turtle species, temperature preference may be influenced by embryonic and post-hatching conditions, such as egg-incubation and acclimation temperature. We tested for effects of embryonic incubation temperature (27.5 °C, 30 °C) and acclimation temperature (20 °C, 25 °C) on the selected temperature and movement patterns of 32 Chrysemys picta bellii (Reptilia: Emydidae) hatchlings in an aquatic thermal gradient of 14-34 °C and in single-temperature (20 °C, 25 °C) control tests. Among 10-11 month old hatchlings, acclimation temperature and egg-incubation temperature influenced temperature selection and movement patterns. Acclimation temperature affected activity and movement: in thermal gradient and single-temperature control tests, 25 °C-acclimated turtles relocated between chambers significantly more frequently than individuals acclimated to 20 °C. Acclimation temperature also affected temperature selection: 20 °C-acclimated turtles selected a specific temperature during gradient tests, but 25 °C-acclimated turtles did not. Among 20 °C-acclimated turtles, egg-incubation temperature was inversely related to selected temperature: hatchling turtles incubated at 27.5 °C selected the warmest temperature available (34 °C); individuals incubated at 30 °C selected the coldest temperature (14 °C). These results suggest that interactions of environmental conditions may influence post-hatching thermoregulatory behavior in C. picta bellii, a factor that ultimately affects fitness.     

Egg components, egg size, and hatchling size in leatherback turtles

Relationships between egg size, egg components, and neonate size have been investigated across a wide range of oviparous taxa. Differences in egg traits among taxa reflect not only phylogenetic differences, but also interactions between biotic (i.e., maternal resource allocation) and abiotic (i.e. nest environment conditions) factors. We examined relationships between egg mass, egg composition, and hatchling size in leatherback turtles (Dermochelys coriacea) because of the unique egg and reproductive characteristics of this species and of sea turtles in general. Albumen comprised 63.0%+/-2.8% (mean+/-S.D.) of egg mass and explained most of the variation in egg mass, whereas yolk comprised only 33.0%+/-2.7%. Additionally, leatherback albumen dry mass was approximately 16% of albumen wet mass. Whereas hatchling mass increased significantly with egg mass (n = 218 clutches), hatchling mass increased by only approximately 2 g for each 10 g increase in egg mass and was approximately 10-20 g greater than yolk mass. Taken together, our results indicate that albumen might play a particularly significant role in leatherback embryonic development, and that leatherback eggs are both capable of water uptake from the nest substrate and also possess a large reservoir of water in the albumen. Relationships between egg mass and egg components, such as variation in egg mass being largely explained by variation in albumen mass and egg mass containing a relatively high proportion of albumen solids, are more similar to bird eggs than to eggs of other non-avian reptiles. However, hatchling mass correlates more with yolk mass than with albumen mass, unlike patterns observed in bird eggs of similar composition.     

Hue discrimination in hatchling loggerhead turtles (Caretta caretta caretta)

Hatchling Atlantic loggerhead turtles (Caretta caretta caretta) were successfully trained to discriminate between broadband hues to obtain food when spatial, intensity, and other confounding cues were randomized with respect to the correct choice. Possible hue preference, the relation of these results to sea-orientation, and the suitability of these animals as subjects in behavioural studies are discussed.     

Roles of thyroid, adrenal and pancreatic hormones on thyroid activity of the soft-shelled turtles Lissemys punctata punctata Bonnoterre

The effects of some exogenous peripheral hormones (thyroxine, corticosterone, epinephrine, norepinephrine and insulin) on thyroid activity were investigated in juvenile female soft-shelled turtles, Lissemys punctata punctata. Each hormone was injected in three different doses (25 microg, 50 microg or 100 microg each per 100 g body weight, once daily at 9 AM) for 10 consecutive days. Thyroid activity was evaluated by gravimetry, histology (epithelial height) and thyroperoxidase assay. The findings revealed that thyroxine in low dose (25 microg) stimulated thyroid activity by increasing the relative thyroid weight, epithelial height and thyroperoxidase activity, but inhibited gland activity at a high dose (100 microg) by decreasing the values of all these parameters. The medium dose (50 microg) had no significant effect. All other hormones, in all doses, significantly decreased thyroid activity by decreasing the values of all the parameters. Thyroid responses to exogenous hormones are generally dose-dependent in turtles. The mechanisms of actions of the hormones administered are suggested.     

Food competition in captive juvenile snapping turtles, Chelydra serpentina

Food-competitive hierarchies and their stability in snapping turtles, Chelydra serpentina, were studied. Nine hatchling turtles were paired against each other in independent tests of food-obtaining dominance. Turtles differed in ability to obtain food. These differences appeared stable over time, and were linear. Neck-stretching, snapping, and clawing were the major behavioural patterns involved in food-appetitive and aggressive encounters.     

The relationship of body size to survivorship of hatchling snapping turtles (Chelydra serpentina): an evaluation of the “bigger is better” hypothesis

In many organisms, body size is positively correlated with traits that are presumably related to fitness. If directional selection frequently favors larger offspring (the “bigger is better” hypothesis), the results of such selection should be detectable with field experiments. We tested the “bigger is better” hypothesis in hatchling snapping turtles (Chelydra serpentina) by conducting one long-term and three short-term experiments on the University of Michigan E.S. George Reserve in southeastern Michigan. In the fall of 1995 and 1996, we released hatchlings at artificial nests separated from the nearest wetland by fences. We recorded the proportion of hatchlings recaptured, the time it took hatchlings to move to fences from artificial nests 45, 55, and 80 m away, and dispersion along the fence. We determined whether the response variables and probability of recapture at fences were associated with hatchling body size. During 1995, average travel times of hatchlings from the experimental nests were not related to distance from the fence; however, time to recapture was positively correlated with dispersion from the zero point on the fence, and the maximum time to reach the fence was almost twice as long for hatchlings from the 80-m nest compared to those from the 45-m nest. Sixty-seven percent of the hatchlings reached the fence and the proportions doing so from each nest were not different. Body size was not significantly related to probability of recapture in either of the 1995 experiments. In 1996, 59% of released hatchlings were recaptured. Time to recapture was not related to dispersion from the zero point or to body size. Cubic spline analysis suggested stabilizing selection on body size. We also conducted a set of long-term hatchling release experiments between 1980–1993 to compare the survival of hatchlings released at nest sites to that of hatchlings released directly into marshes, and we looked for relationships between survivorship and hatchling body size. During 7 years in which more than 30 hatchlings were released, 413 hatchlings were released directly into the marsh and 262 were released at nests: their probability of survival did not differ. Over all years, for both release groups combined and for each group separately, survival was not related to body size. In 1983 alone, survival was also not related to body size for either group or for both groups combined. In our three short-term experiments and one long-term experiment, we found no evidence to support the “bigger is better” hypothesis. When selection on body size did occur, selection was stabilizing, not directional for larger size. Received: 4 June 1998 / Accepted: 24 June 1999