Summary 1. Freshwater fishes are the most northerly of freshwater ectotherms, followed by frogs. North American freshwater snakes, turtles, and salamanders do not range farther north than southernmost Canada. 2. Freezing and desiccation are the main challenges during terrestrial hibernation of ectotherms. Oxygen depletion, water balance, and ionic balance are the major problems for air breathing ectotherms that hibernate underwater. 3. The importance of accumulation of energy stores for overwintering among fishes depends upon the length and severity of the winters, whether or not there is springtime reproduction, body size, latitude, and the availability and use of food during overwintering. 4. Fishes can decrease energy demands during the winter by reductions in activity, metabolic depression, and entrance in semi‐torpidity. 5. Adaptations for coping with hypoxia and anoxia among overwintering freshwater fishes may include metabolic depression, a decrease in blood O2 affinity, microhabitat selection, air breathing, short‐distance migration, biochemical modifications aimed at adjusting glycolytic rates, and alcoholic fermentation. 6. Freshwater turtles have a worldwide northern limit of approximately 50° N, which means that some species spend about half of their lives hibernating. 7. Aquatic turtles normally hibernate underwater, although occasionally they hibernate on land. In water they usually hibernate in a hypoxic or anoxic (mud) environment and in relatively shallow water. Wintertime movements of unknown frequency occur in some species. 8. The hatchlings of many turtle species can overwinter in the nest. Among northern species this behaviour is most common among painted turtles, whose hatchlings can withstand freezing. 9. Mortality among adult turtles is probably highest during the hibernation cycle. 10. Temperature appears to the most important cue for entry and exit from hibernation among freshwater turtles. 11. Little is known of the energetics of overwintering turtles. Energy stores for overwintering may be more important at lower latitudes than at higher ones, due to the higher metabolic rates of overwintering, but non‐feeding, southern turtles. 12. The ability of turtles to tolerate submergence is a function of temperature, degree of water oxygenation, latitude of origin, efficacy of extrapulmonary respiratory pathways, and metabolic rate. 13. For turtles that hibernate in an anoxic hibernaculum, and for those without sufficient extrapulmonary uptake of O2 to allow metabolism to be completely aerobic, the most important physiological perturbation is an acidosis developed from a continuing production of lactate. If sufficient O2 can be obtained, the most likely factors limiting hibernation time are water balance and ion balance. 14. Mechanisms of turtles for coping with acidosis include metabolic depression, integumental CO2 loss, bicarbonate buffering, and changes in ion concentrations that minimize the decrease in SID (strong ion difference). The most important among the latter are a decrease in plas...

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Turtles are a small taxon that has nevertheless attracted much attention from biologists for centuries. However, a major portion of their life cycle has received relatively little attention until recently - namely what turtles are doing, and how they are doing it, during the winter. In the northern parts of their ranges in North America, turtles may spend more than half of their lives in an overwintering state. In this review, I emphasise the ecological aspects of overwintering among turtles, and consider how overwintering stresses affect the physiology, behaviour, distributions, and life histories of various species. Sea turtles are the only group of turtles that migrate extensively, and can therefore avoid northern winters. Nevertheless, each year a number of turtles, largely juveniles, are killed when trapped by cold fronts before they move to safer waters. Evidently this risk is an acceptable trade-off for the benefits to a population of inhabiting northern developmental habitats during the summer. Terrestrial turtles pass the winter underground, either in burrows that they excavate or that are preformed. These refugia must provide protection against desiccation and lethal freezing levels. Some burrows are extensive (tortoise genus Gopherus), while others are shallow, or the turtles may simply dig into the ground to a safe depth (turtle genus Terrapene). In the latter genus, freeze tolerance may play an adaptive role. Most non-marine aquatic turtles overwinter underwater, although Clemmys (Actinemys) marmorata routinely overwinters on land when it occurs in riverine habitats, Kinosternon subrubrum often overwinters on land, and several others may overwinter terrestrially on occasion, especially in more southern climates. For northern species that overwinter underwater, there are two physiological groupings, those that are anoxia-tolerant and those that are relatively anoxia-intolerant. All species fare well physiologically in water with a high partial pressure of oxygen (PO2). A lack of anoxia tolerance limits the types of habitats that a freshwater turtle may live in, since unlike sea turtles, they cannot travel long distances to hibernate. Hatchlings of some species of turtles spend their first winter in or below the nest cavity, while hatchlings of other species in the same area, including northern areas, emerge in the autumn and presumably hibernate underwater. All hatchlings are relatively anoxia-intolerant, and there are no studies to date of where hatchling turtles that do not overwinter in or below the nest cavity spend their first winter. Equally little is known of the ontogeny of anoxia tolerance, other than that adults of all species are more anoxia-tolerant than their hatchlings, probably because of their better ossified shells, which provide adults with more buffer reserves and a larger site in which to sequester lactate. The northern limits of turtles are most likely determined by reproductive limitations (time for egg-laying, incubation, and hatching) than by the rigors of hibernation. Mortalit...

In cold-temperate climates, overwintering aquatic ranid frogs must survive prolonged periods of low temperature, often accompanied by low levels of dissolved oxygen. They must do so with the energy stores acquired prior to the onset of winter. Overwintering mortality is a significant factor in their life history, occasionally reaching 100% due to freezing and/or anoxia. Many species of northern ranid frogs overwinter in the tadpole stage, which increases survival during hypoxic episodes relative to adults, as well as allowing for larger sizes at metamorphosis. At temperatures below 5 degrees C, submerged ranid frogs are capable of acquiring adequate oxygen via cutaneous gas exchange over a wide range of ambient oxygen partial pressures (PO(2)), and possess numerous physiological and behavioural mechanisms that allow them to maintain normal rates of oxygen uptake across the skin at a relatively low PO(2). At levels of oxygen near and below the critical PO(2) that allows for aerobic metabolism, frogs must adopt biochemical mechanisms that act to minimise oxygen utilisation and assist in maintaining an aerobic state to survive overwintering. These mechanisms include alterations in mitochondrial metabolism and affinity, changes in membrane permeability, alterations in water balance, and reduction in cellular electrochemical gradients, all of which lead to an overall reduction in whole-animal metabolism. Winter energetic requirements are fueled by the energy stores in liver, muscle, and fat depots, which are likely to be sufficient when the water is cold and well oxygenated. However, under hypoxic conditions fat stores cannot be utilised efficiently and glycogen stores are used up rapidly due to recruitment of anaerobiosis. Since ranid frogs have minimal tolerance to anoxia, it is untenable to suggest that they spend a significant portion of the winter buried in anoxic mud, but instead utilise a suite of behavioural and physiological mechanisms geared to optimal survival in cold, hypoxic conditions.

Successful overwintering under ice by an air-breathing vertebrate requires either effective aquatic respiration if dissolved O 2 is available or the capacity for prolonged anaerobic metabolism if O 2 supplies are limiting. Frogs can remain aerobic for many weeks when submerged at low temperature, even at water PO 2 as low as 30 mmHg, but are unable to survive even 1 week in anoxic water. Fuel reserves of hibernating frogs limit aerobic submergence, whereas acidosis may limit anoxic submergence. Freshwater turtles can also satisfy all or most of their O 2 needs in wellaerated water at low temperature by aquatic respiration, but certain species, in particular painted and snapping turtles, can also survive for up to 4-5 months without O 2 . Key adaptations of the painted turtles, and presumably snapping turtles, include metabolic depression and the exploitation of the shell and other bones to buffer lactic acid. As in frogs, glycogen and glucose are the only fuel sources during anoxia, and stores do not seem to be limiting in the painted turtle. Significant differences in anoxia tolerance exist among chelonian species that can be attributed, at least in part, to the magnitude of metabolic depression, the effectiveness of lactic acid buffering, and the size of glycogen stores.

Abstract. The painted turtle, Chrysemys picta, is currently recognized as a continentally distributed polytypic species, ranging across North America from southern Canada to extreme northern Mexico. We analyzed variation in the rapidly evolving mitochondrial control region (CR) in 241 turtles from 117 localities across this range to examine whether the painted turtle represents a continentally distributed species based on molecular analysis. We found strong support for the novel hypothesis that C. p. dorsalis is the sister group to all remaining Chrysemys, with the remaining Chrysemys falling into a single, extremely wide-ranging and genetically undifferentiated species. Given our goal of an evolutionarily accurate taxonomy, we propose that two evolutionary lineages be recognized as species within Chrysemys: C. dorsalis (Agassiz 1857) in the southern Mississippi drainage region, and C. picta (Schneider 1783) from the rest of the range of the genus. Neither molecular nor recent morphological analyses argue for the hybrid origin of C. p. marginata as previously proposed. Within C. picta, we find evidence of at least two independent range expansions into previously glaciated regions of North America, one into New England and the other into the upper Midwest. We further find evidence of a massive extinction/recolonization event across the Great Plains/Rocky Mountain region encompassing over half the continental United States. The timing and extent of this colonization is consistent with a recently proposed regional aridification as the Laurentide ice sheets receded approximately 14,000 years ago, and we tentatively propose this paleoclimatological event as a major factor shaping genetic variation in Chrysemys.

The painted turtle, Chrysemys picta, is currently recognized as a continentally distributed polytypic species, ranging across North America from southern Canada to extreme northern Mexico. We analyzed variation in the rapidly evolving mitochondrial control region (CR) in 241 turtles from 117 localities across this range to examine whether the painted turtle represents a continentally distributed species based on molecular analysis. We found strong support for the novel hypothesis that C. p. dorsalis is the sister group to all remaining Chrysemys, with the remaining Chrysemys falling into a single, extremely wide-ranging and genetically undifferentiated species. Given our goal of an evolutionarily accurate taxonomy, we propose that two evolutionary lineages be recognized as species within Chrysemys: C. dorsalis (Agassiz 1857) in the southern Mississippi drainage region, and C. picta (Schneider 1783) from the rest of the range of the genus. Neither molecular nor recent morphological analyses argue for the hybrid origin of C. p. marginata as previously proposed. Within C. picta, we find evidence of at least two independent range expansions into previously glaciated regions of North America, one into New England and the other into the upper Midwest. We further find evidence of a massive extinction/recolonization event across the Great Plains/Rocky Mountain region encompassing over half the continental United States. The timing and extent of this colonization is consistent with a recently proposed regional aridification as the Laurentide ice sheets receded approximately 14,000 years ago, and we tentatively propose this paleoclimatological event as a major factor shaping genetic variation in Chrysemys.

To evaluate shell and bone buffering of lactic acid during acidosis at 3 degrees C, turtles were submerged in anoxic or aerated water and tested at intervals for blood acid-base status and plasma ions and for bone and shell percent water, percent ash, and concentrations of lactate, Ca(2+), Mg(2+), P(i), Na(+), and K(+). After 125 days, plasma lactate concentration rose from 1.6 +/- 0.2 mM (mean +/- SE) to 155.2 +/- 10.8 mM in the anoxic group but only to 25.2 +/- 6.4 mM in the aerated group. The acid-base state of the normoxic animals was stable after 25 days of submergence. Plasma calcium concentration (¿Ca(2+)) rose during anoxia from 3.2 +/- 0.2 to 46.0 +/- 0.6 mM and ¿Mg(2+) from 2.7 +/- 0.2 to 12.2 +/- 0.6 mM. Both shell and bone accumulated lactate to concentrations of 135.6 +/- 35.2 and 163.6 +/- 5.1 mmol/kg wet wt, respectively, after 125 days anoxia. Shell and bone ¿Na(+) both fell during anoxia but the fate of this Na(+) is uncertain because plasma ¿Na(+) also fell. No other shell ions changed significantly in concentration, although the concentrations of both bone calcium and bone potassium changed significantly. Control shell water (27.8 +/- 0.6%) was less than bone water (33.6 +/- 1.1%), but neither changed during submergence. Shell ash (44.7 +/- 0.8%) remained unchanged, but bone ash (41.0 +/- 1.0%) fell significantly. We conclude that bone, as well as shell, accumulate lactate when plasma lactate is elevated, and that both export sodium carbonate, as well as calcium and magnesium carbonates, to supplement ECF buffering.