Energetic costs of bill heat exchange demonstrate contributions to thermoregulation at high temperatures in toco toucans (Ramphastos toco).

Body temperature regulation under changes in ambient temperature involves adjustments in heat production and heat exchange rates between the animal and the environment. One mechanism involves the modulation of the surface temperature of specific areas of the body through vasomotor adjustment. In homeotherms, this thermoregulatory adjustment is essential for the maintenance of body temperature over a moderate temperature range, known as the thermal neutral zone (TNZ). The bill of the toco toucan (Ramphastos toco) has been described as a highly efficient thermal window and hypothesized to assist in the thermal homeostasis of this bird. Herein, we directly evaluated the contribution of heat exchange through the bill of the toco toucan and role of the bill in the delimitation of the TNZ. To do this, we measured metabolic rate (MR), via oxygen consumption, over a range of ambient temperatures from 0 to 35°C. MR measurements were made in birds with the bill intact and with the bill insulated. The limits of the TNZ did not differ between treatments, ranging from 10.8 to 25.0°C. The MR differed among treatments only at elevated temperatures (30 and 35°C), reaching 0.92±0.11 ml O2 g-1 h-1 (mean±s.d.) for the intact group and 1.13±0.13 ml O2 g-1 h-1 for the insulated group. These results indicate that although heat dissipation through the bill does not contribute significantly to widening of the TNZ, it may well be critically important in assisting body temperature regulation at higher temperatures extending above the upper limit of the TNZ.

Empirical estimation of skin resistance to water loss in amphibians: agar evaluation as a non-resistance model to evaporation.

Total resistance (rt) to evaporative water loss (EWL) in amphibians is given by the sum of the boundary layer (rb) and the skin resistance (rs). Thus, rs can be determined if the rb component is defined (rs=rt-rb). The use of agar models has become the standard technique to estimate rb under the assumption that the agar surface imposes no barrier to evaporation (rs=0). We evaluated this assumption by determining EWL rates and rb values from exposed surfaces of free water, a physiological solution mimicking the osmotic properties of a generalized amphibian, and agar gels prepared at various concentrations using either water or physiological solution as diluent. Water evaporation was affected by both the presence of solutes and agar concentration. Models prepared with agar at 5% concentration in water provided the most practical and appropriate proxy for the estimation of rb.

Maintained barostatic regulation of heart rate in digesting snakes (Boa constrictor).

When snakes digest large meals, heart rate is accelerated by withdrawal of vagal tone and an increased non-adrenergic-non-cholinergic tone that seems to stem from circulating blood-borne factors exerting positive chronotropic effects. To investigate whether this tonic elevation of heart rate impairs the ability for autonomic regulation of heart during digestion, we characterised heart rate responses to pharmacological manipulation of blood pressure in the snake Boa constrictor through serial injections of sodium nitroprusside and phenylephrine. Both fasting and digesting snakes responded with a robust tachycardia to hypotension induced by sodium nitroprusside, with digesting snakes attaining higher maximal heart rates than fasting snakes. Both fasting and digesting snakes exhibited small reductions of the cardiac chronotropic response to hypertension, induced by injection of phenylephrine. All heart rate changes were abolished by autonomic blockade with the combination of atropine and propranolol. The digesting snakes retained the capacity for compensatory heart rate responses to hypotension, despite their higher resting values, and the upward shift of the barostatic response curve enables snakes to maintain the cardiac limb of barostatic regulation for blood pressure regulation.

Feeding alters the preferred body temperature of Cururu toads, Rhinella diptycha (Anura, Bufonidae).

Ectothermic organisms depend primarily on external heat sources and behavioural adjustments to regulate body temperature. Under controlled conditions, in a thermal gradient, body temperature often clusters around a more or less defined range of preferred body temperatures (Tpref). However, Tpref may be modified in response to environmental parameters and/or physiological state. For example, meal ingestion is sometimes followed by a post-prandial thermophilic response leading to a transient increment in Tpref. Although thought to optimize digestive processes, its occurrence, magnitude, and possible determinants remains scarcely documented for anuran amphibians. Herein, we investigated whether the Cururu toad, Rhinella diptycha, exhibits a post-prandial thermophilic response by monitoring the body temperature of fasting and fed toads while they were maintained in a thermal gradient. We found that the toads' Tpref increased by about 13% from day 2 to 4 after feeding, in comparison with the Tpref recorded under fasting. Also, fed animals exhibited a broader range for Tpref at days 2 and 3 post-prandial, which reflects a greater level of locomotor activity compared to fasting individuals. We conclude that R. diptycha is capable to exhibit a post-prandial thermophilic response under the controlled conditions of a thermal gradient. Although this thermoregulatory adjustment is thought to optimize meal digestion yielding important energetic and ecological benefits, its occurrence in anuran amphibians in nature remains uncertain.

Effects of temperature and meal size on the postprandial metabolic response of Leptodactylus latrans (Anura, Leptodactylidae).

The postprandial increment of metabolism, often referred to as specific dynamic action (SDA), encompasses the summed costs of meal ingestion, digestion, absorption, and assimilation. Different SDA parameters, such as its magnitude, duration, and relative cost, are affected by a diverse set of environmental and physiological determinants, including meal size and body temperature. While the influence of these variables has been thoroughly examined in most ectothermic vertebrate groups, few studies have focused on the determinants and consequences of the SDA response in anuran amphibians. Thus, we examined the effects of meal size and body temperature on the SDA response of a Neotropical frog, Leptodactylus latrans, by measuring the rates of oxygen consumption of frogs while fasting and after being fed meals of different sizes at different temperatures. SDA lasted from 3 to 5 days and increased with meal size for frogs fed meals equivalent to 5-15% of their body mass. SDA cost was not affected by meal size and averaged 14.6% of the caloric content of the meal. Temperature increment was accompanied by proportional increases in metabolic rate and shorter SDA duration. The relative SDA cost averaged 14.8% of the meal caloric content at 20°C and 25°C, but increased to 23.5% at 30°C. Our results indicate that meal size imposes no physiological or energetic constraint to L. latrans feeding. Digestion at temperatures near the thermal preference of the species seems to optimize energetic return, whereas the shortening of SDA duration at higher temperatures may provide significant ecological advantages.

Exposure to fluctuating temperatures leads to reduced immunity and to stress response in rattlesnakes.

Ectothermic organisms often experience considerable variation in their body temperature throughout the circadian cycle. However, studies focusing on the measurement of physiological traits are usually performed under constant temperature regimes. This mismatch between thermal exposure in the field and experimental conditions could act as a stressor agent, as physiological functions are strongly influenced by temperature. Herein, we asked the question whether constant thermal regimes would cause a stress response and impact the immunity of the South American rattlesnake (Crotalus durissus) when compared with a fluctuating thermal regime. We addressed this question by determining heterophil:lymphocyte (H:L) ratio, plasma bacteria-killing ability (BKA) and corticosterone (CORT) levels in snakes kept under a constant temperature regime at 30°C, and under a fluctuating regime that oscillated between 25°C at night and 35°C during the day. The experiments had a mirrored design, in which half of the snakes were subjected to a fluctuating-to-constant treatment, while the other half was exposed to a constant-to-fluctuating treatment. The shift from constant to fluctuating thermal regime was accompanied by an increase in plasma CORT levels, indicating the activation of a stress response. Exposure to a fluctuating thermal regime at the onset of the experiments induced a decrease in the BKA of rattlesnakes. H:L ratio was not affected by treatments and, therefore, the shift between thermal regimes seems to have acted as a low-intensity stressor. Our results suggest that removal from temperatures close to the snake's preferred body temperature triggers a stress response in rattlesnakes.

Biophysical Modeling of Water Economy Can Explain Geographic Gradient of Body Size in Anurans.

Geographical gradients of body size express climate-driven constraints on animals, but whether they exist and what causes them in ectotherms remains contentious. For amphibians, the water conservation hypothesis posits that larger bodies reduce evaporative water loss (EWL) along dehydrating gradients. To address this hypothesis mechanistically, we build on well-established biophysical equations of water exchange in anurans to propose a state-transition model that predicts an increase of either body size or resistance to EWL as alternative specialization along dehydrating gradients. The model predicts that species whose water economy is more sensitive to variation in body size than to variation in resistance to EWL should increase in size in response to increasing potential evapotranspiration (PET). To evaluate the model predictions, we combine physiological measurements of resistance to EWL with geographic data of body size for four different anuran species. Only one species, Dendropsophus minutus, was predicted to exhibit a positive body size-PET relationship. Results were as predicted for all cases, with one species-Boana faber-showing a negative relationship. Based on an empirically verified mathematical model, we show that clines of body size among anurans depend on the current values of those traits and emerge as an advantage for water conservation. Our model offers a mechanistic and compelling explanation for the cause and variation of gradients of body size in anurans.

Droplet bubbling evaporatively cools a blowfly.

Terrestrial animals often use evaporative cooling to lower body temperature. Evaporation can occur from humid body surfaces or from fluids interfaced to the environment through a number of different mechanisms, such as sweating or panting. In Diptera, some flies move tidally a droplet of fluid out and then back in the buccopharyngeal cavity for a repeated number of cycles before eventually ingesting it. This is referred to as the bubbling behaviour. The droplet fluid consists of a mix of liquids from the ingested food, enzymes from the salivary glands, and antimicrobials, associated to the crop organ system, with evidence pointing to a role in liquid meal dehydration. Herein, we demonstrate that the bubbling behaviour also serves as an effective thermoregulatory mechanism to lower body temperature by means of evaporative cooling. In the blowfly, Chrysomya megacephala, infrared imaging revealed that as the droplet is extruded, evaporation lowers the fluid´s temperature, which, upon its re-ingestion, lowers the blowfly's body temperature. This effect is most prominent at the cephalic region, less in the thorax, and then in the abdomen. Bubbling frequency increases with ambient temperature, while its cooling efficiency decreases at high air humidities. Heat transfer calculations show that droplet cooling depends on a special heat-exchange dynamic, which result in the exponential activation of the cooling effect.

The consequences of seasonal fasting during the dormancy of tegu lizards (Salvator merianae) on their postprandial metabolic response.

Tegu lizards (Salvator merianae) aestivate for up to 5 months during Brazil's winter, when they retreat to burrows and halt most activities. Dormant tegus reduce their gastrointestinal (GI) mass, which allows a substantial energy economy. This strategy, however, implies that the first post-dormancy digestion would be more costly than subsequent feeding episodes as a result of GI atrophy. To address this, we determined the postprandial metabolic response (SDA) of the first (M1), second (M2) and several (RM) feeding episodes after tegus' dormancy. Another group of tegus (PF) was subjected to an extra 50 day fasting period after arousal. Glucose, triglycerides and uric acid levels were checked before and after feeding. M1 digestion lasted twice as long and cost twofold more when compared with M2 or RM, in agreement with the idea that GI atrophy inflates digestion cost at the first post-dormancy meal. The SDA response was similar in M2 and RM, suggesting that the GI tract was fully reorganized after the first feeding. The SDA cost was equal in PF and RM, implying that the change in state per se (dormant to arousal) triggers the regrowth of GI, independently of feeding. Fasting tegus at M1 presented higher triglyceride and lower uric acid levels than fed tegus, indicating that fasting is mainly sustained by fat storage. Our results show that seasonal fasting imposes an extra digestion cost to tegus following their next feeding, which is fully paid during their first digestion. This surplus cost, however, is negligible compared with the overall energetic savings from GI tract atrophy during the dormancy period.

Trading heat and hops for water: Dehydration effects on locomotor performance, thermal limits, and thermoregulatory behavior of a terrestrial toad.

Due to their highly permeable skin and ectothermy, terrestrial amphibians are challenged by compromises between water balance and body temperature regulation. The way in which such compromises are accommodated, under a range of temperatures and dehydration levels, impacts importantly the behavior and ecology of amphibians. Thus, using the terrestrial toad Rhinella schneideri as a model organism, the goals of this study were twofold. First, we determined how the thermal sensitivity of a centrally relevant trait-locomotion-was affected by dehydration. Secondly, we examined the effects of the same levels of dehydration on thermal preference and thermal tolerance. As dehydration becomes more severe, the optimal temperature for locomotor performance was lowered and performance breadth narrower. Similarly, dehydration was accompanied by a decrease in the thermal tolerance range. Such a decrease was caused by both an increase in the critical minimal temperature and a decrease in the thermal maximal temperature, with the latter changing more markedly. In general, our results show that the negative effects of dehydration on behavioral performance and thermal tolerance are, at least partially, counteracted by concurrent adjustments in thermal preference. We discuss some of the potential implications of this observation for the conservation of anuran amphibians.

Not Good, but Not All Bad: Dehydration Effects on Body Fluids, Organ Masses, and Water Flux through the Skin of Rhinella schneideri (Amphibia, Bufonidae).

Because of their permeable skin, terrestrial amphibians are constantly challenged by the potential risk of dehydration. However, some of the physiological consequences associated with dehydration may affect aspects that are themselves relevant to the regulation of water balance. Accordingly, we examined the effects of graded levels of dehydration on the rates of evaporative water loss and water absorption through the skin in the terrestrial Neotropical toad, Rhinella schneideri. Concomitantly, we monitored the effects of dehydration on the mass of visceral organs; hematocrit and hemoglobin content; plasma osmolality; and plasma concentration of urea, sodium, chloride, and potassium. We found that dehydration caused an increase in the concentration of body fluids, as indicated by virtually all the parameters examined. There was a proportional change in the relative masses of visceral organs, except for the liver and kidneys, which exhibited a decrease in their relative masses greater than the whole-body level of dehydration. Changes-or the preservation-of relative organ masses during dehydration may be explained by organ-specific physiological adjustments in response to the functional stress introduced by the dehydration itself. As dehydration progressed, evaporative water loss diminished and water reabsorption increased. In both cases, the increase in body fluid concentration associated with the dehydration provided the osmotic driver for these changes in water flux. Additionally, dehydration-induced alterations on the cutaneous barrier may also have contributed to the decrease in water flux. Dehydration, therefore, while posing a considerable challenge on the water balance regulation of anurans, paradoxically facilitates water conservation and absorption.

The evolution of the meatal chamber in crocodyliforms.

The unique outer ear of crocodylians consists of a large meatal chamber (MC) concealed by a pair of muscular earlids that shape a large part of the animal's head. This chamber is limited medially by the enlarged tympanic membrane. Yet, the anatomy of this distinctive and complex region is underexplored and its evolutionary history untraced. The osteology and soft tissues of the MC in extant crocodylians was analysed to describe it and establish osteological correlates within this region. A broad survey of the osteological correlates was conducted in major clades of fossil crocodyliforms to estimate evolutionary trends of the MC. The reorganization of the MC at the origin of crocodyliforms includes characters also present in more basal taxa such as 'sphenosuchians' as well as unique traits of crocodyliforms. Three major patterns are recognized in the MC of basal mesoeucrocodylians. The distinct 'thalattosuchian pattern' indicates that extensive modifications occurred in this clade of aquatic fossil crocodyliforms, even when multiple alternative phylogenetic positions are taken into account. Some traits already established in putative closely related clades are absent or modified in this group. The 'basal notosuchian/sebecian pattern' is widespread among basal metasuchians, and establishes for the first time characters maintained later in neosuchians and extant forms. The 'advanced notosuchian pattern' includes modifications of the MC possibly related to a terrestrial lifestyle and potentially a structure analogous to the mammalian pinna. The main variation in the MC of neosuchians is associated with the homoplastic secondary opening of the cranioquadrate passage. The inferred phylogenetic trends in the crocodyliform MC suggest the great anatomical disparity in this region followed a complex evolutionary pattern, and tympanic hearing played an important role in the origin and diversification of Crocodyliformes.

Seasonal reproductive endothermy in tegu lizards.

With some notable exceptions, small ectothermic vertebrates are incapable of endogenously sustaining a body temperature substantially above ambient temperature. This view was challenged by our observations of nighttime body temperatures sustained well above ambient (up to 10°C) during the reproductive season in tegu lizards (~2 kg). This led us to hypothesize that tegus have an enhanced capacity to augment heat production and heat conservation. Increased metabolic rates and decreased thermal conductance are the same mechanisms involved in body temperature regulation in those vertebrates traditionally acknowledged as "true endotherms": the birds and mammals. The appreciation that a modern ectotherm the size of the earliest mammals can sustain an elevated body temperature through metabolic rates approaching that of endotherms enlightens the debate over endothermy origins, providing support for the parental care model of endothermy, but not for the assimilation capacity model of endothermy. It also indicates that, contrary to prevailing notions, ectotherms can engage in facultative endothermy, providing a physiological analog in the evolutionary transition to true endothermy.

Physiological responses of Brazilian amphibians to an enzootic infection of the chytrid fungus Batrachochytrium dendrobatidis.

Pathophysiological effects of clinical chytridiomycosis in amphibians include disorders of cutaneous osmoregulation and disruption of the ability to rehydrate, which can lead to decreased host fitness or mortality. Less attention has been given to physiological responses of hosts where enzootic infections of Batrachochytrium dendrobatidis (Bd) do not cause apparent population declines in the wild. Here, we experimentally tested whether an enzootic strain of Bd causes significant mortality and alters host water balance (evaporative water loss, EWL; skin resistance, R(s); and water uptake, WU) in individuals of 3 Brazilian amphibian species (Dendropsophus minutus, n = 19; Ischnocnema parva, n = 17; Brachycephalus pitanga, n = 15). Infections with enzootic Bd caused no significant mortality, but we found an increase in R(s) in 1 host species concomitant with a reduction in EWL. These results suggest that enzootic Bd infections can indeed cause sub-lethal effects that could lead to reduction of host fitness in Brazilian frogs and that these effects vary among species. Thus, our findings underscore the need for further assessment of physiological responses to Bd infections in different host species, even in cases of sub-clinical chytridiomycosis and long-term enzootic infections in natural populations.

Winter metabolic depression does not change arterial baroreflex control of heart rate in the tegu lizard Salvator merianae.

Baroreflex regulation of blood pressure is important for maintaining appropriate tissue perfusion. Although temperature affects heart rate (fH) reflex regulation in some reptiles and toads, no data are available on the influence of temperature-independent metabolic states on baroreflex. The South American tegu lizard Salvator merianae exhibits a clear seasonal cycle of activity decreasing fH along with winter metabolic downregulation, independent of body temperature. Through pharmacological interventions (phenylephrine and sodium nitroprusside), the baroreflex control of fH was studied at ∼ 25 °C in spring-summer- and winter-acclimated tegus. In winter lizards, resting and minimum fH were lower than in spring-summer animals (respectively, 13.3 ± 0.82 versus 10.3 ± 0.81 and 11.2 ± 0.65 versus 7.97 ± 0.88 beats min(-1)), while no acclimation differences occurred in resting blood pressure (5.14 ± 0.38 versus 5.06 ± 0.56 kPa), baroreflex gain (94.3 ± 10.7 versus 138.7 ± 30.3% kPa(-1)) or rate-pressure product (an index of myocardial activity). Vagal tone exceeded the sympathetic tone of fH, especially in the winter group. Therefore, despite the lower fH, winter acclimation does not diminish the fH baroreflex responses or rate-pressure product, possibly because of increased stroke volume that may arise because of heart hypertrophy. Independent of acclimation, fH responded more to hypotension than to hypertension. This should imply that tegus, which have no pressure separation within the single heart ventricle, must have other protection mechanisms against pulmonary hypertension or oedema, presumably through lymphatic drainage and/or vagal vasoconstriction of pulmonary artery. Such a predominant fH reflex response to hypotension, previously observed in anurans, crocodilians and mammals, may be a common feature of tetrapods.

Daily and annual cycles in thermoregulatory behaviour and cardio-respiratory physiology of black and white tegu lizards.

This study was designed to determine the manner in which metabolism is suppressed during dormancy in black and white tegu lizards (Tupinambis merianae). To this end, heart rate (fH), respiration rate (fR), and deep body temperature (Tb) were continuously monitored in outdoor enclosures by radio-telemetry for nine months. There was a continuous decline in nighttime breathing and heart rate, at constant Tb, throughout the late summer and fall suggestive of an active metabolic suppression that developed progressively at night preceding the entrance into dormancy. During the day, however, the tegus still emerged to bask. In May, when the tegus made a behavioural commitment to dormancy, Tb (day and night) fell to match burrow temperature, accompanied by a further reduction in fH and fR. Tegus, under the conditions of this study, did arouse periodically during dormancy. There was a complex interplay between changes in fH and Tb associated with the direct effects of temperature and the indirect effects of thermoregulation, activity, and changes in metabolism. This interplay gave rise to a daily hysteresis in the fH/Tb relationship reflective of the physiological changes associated with warming and cooling as preferred Tb alternated between daytime and nighttime levels. The shape of the hysteresis curve varied with season along with changes in metabolic state and daytime and nighttime body temperature preferences.

The relationship between body temperature, heart rate, breathing rate, and rate of oxygen consumption, in the tegu lizard (Tupinambis merianae) at various levels of activity.

The present study determined whether EEG and/or EMG recordings could be used to reliably define activity states in the Brazilian black and white tegu lizard (Tupinambis merianae) and then examined the interactive effects of temperature and activity states on strategies for matching O2 supply and demand. In a first series of experiments, the rate of oxygen consumption (VO2), breathing frequency (fR), heart rate (fH), and EEG and EMG (neck muscle) activity were measured in different sleep/wake states (sleeping, awake but quiet, alert, or moving). In general, metabolic and cardio-respiratory changes were better indictors of the transition from sleep to wake than were changes in the EEG and EMG. In a second series of experiments, the interactive effects of temperature (17, 27 and 37 °C) and activity states on fR, tidal volume (VT), the fraction of oxygen extracted from the lung per breath (FIO2-FEO2), fH, and the cardiac O2 pulse were quantified to determine the relative roles of each of these variables in accommodating changes in VO2. The increases in oxygen supply to meet temperature- and activity-induced increases in oxygen demand were produced almost exclusively by increases in fH and fR. Regression analysis showed that the effects of temperature and activity state on the relationships between fH, fR and VO2 was to extend a common relationship along a single curve, rather than separate relationships for each metabolic state. For these lizards, the predictive powers of fR and fH were maximized when the effects of changes in temperature, digestive state and activity were pooled. However, the best r(2) values obtained were 0.63 and 0.74 using fR and fH as predictors of metabolic rate, respectively.

Anuran skin and basking behavior: The case of the treefrog Bokermannohyla alvarengai (Bokermann, 1956).

We investigated the morphology of the skin and the biochemistry of the lipids in the skin secretion of Bokermannohyla alvarengai, a montane treefrog that is known to bask regularly, motionless in full sunlight for extended periods of time. Our primary goal was to identify structural and biochemical modifications that might assist this frog species to accommodate the conflicting demands for heat exchange and water balance while basking. The modulation of heat exchange in basking B. alvarengai involves changes in skin coloration. We found that this response was supported by a prominent monolayer of large iridophores, whose light reflectance property is adjusted by the response of intervening melanophores. Mucosubstances and lipid compounds, mainly consisted of saturated fatty acids and presumably secreted from granular glands, were detected on the skin of B. alvarengai. These compounds formed an extra-epidermal layer over the animal's dorsal surface that might assist in the prevention of excessive water loss through evaporation. Additionally, we found well-developed skin folds at the ventral region of the frogs that lead to an increment of surface area. This feature combined with the extensive hypervascularization, also noticed for the skin of B. alvarengai, may play an important role in water reabsorption. The suite of structural and biochemical modifications identified for the integument of B. alvarengai seems to conjugate aspects relevant to both, heat exchange and water balance, allowing for this species to explore basking as an efficient thermoregulatory strategy.

Blood oxygen affinity increases during digestion in the South American rattlesnake, Crotalus durissus terrificus.

Digesting snakes experience massive increases in metabolism that can last for many days and are accompanied by adjustments in the oxygen transport cascade. Accordingly, we examined the oxygen-binding properties of the blood in the South American rattlesnake (Crotalus durissus terrificus) during fasting and 24 and 48h after the snakes have ingested a rodent meal corresponding to 15% (±2%) of its own body mass. In general, oxygen-hemoglobin (Hb-O2) affinity was significantly increased 24h post-feeding, and then returned toward fasting values within 48h post-feeding. Content of organic phosphates ([NTP] and [NTP]/[Hb]), hemoglobin cooperativity (Hill's n), and Bohr Effect (ΔlogP50/ΔpH) were not affected by feeding. The postprandial increase in Hb-O2 affinity in the South American rattlesnake can be almost entirely ascribed by the moderate alkaline tide that follows meal ingestion. In general, digesting snakes were able to regulate blood metabolites at quite constant levels (e.g., plasma osmolality, lactate, glucose, and total protein levels). The level of circulating lipids, however, was considerably increased, which may be related to their mobilization, since lipids are known to be incorporated by the enterocytes after snakes have fed. In conclusion, our results indicate that the exceptional metabolic increment exhibited by C. d. terrificus during meal digestion is entirely supported by the aerobic pathways and that among the attending cardiorespiratory adjustments, pulmonary Hb-O2 loading is likely improved due to the increment in blood O2 affinity.

Thermal biology of the toad Rhinella schneideri in a seminatural environment in southeastern Brazil.

The toad, Rhinella schneideri, is a large-bodied anuran amphibian with a broad distribution over South America. R. schneideri is known to be active at night during the warm/rainy months and goes into estivation during the dry/cold months; however, there is no data on the range of body temperatures (Tb) experienced by this toad in the field, and how environmental factors, thermoregulatory behaviors or activity influence them. By using implantable temperature dataloggers, we provide an examination of Tb variation during an entire year under a seminatural setting (emulating its natural habitat) monitored with thermosensors. We also used data on preferred Tb, allowing us to express the effectiveness of thermoregulation quantitatively. Paralleling its cycle of activity, R. schneideri exhibited differences in its daily and seasonal profile of Tb variation. During the active season, toads spent daytime hours in shelters and, therefore, did not explore microhabitats with higher thermal quality, such as open areas in the sun. At nighttime, the thermal suitability of microhabitats shifted as exposed microhabitats experienced greater temperature drops than the more insulated shelter. As toads became active at night, they were driven to the more exposed areas and, as a result, thermoregulatory effectiveness decreased. Our results, therefore, indicate that, during the active season, a compromise between thermoregulation and nocturnal activity may be at play. During the estivation period, R. schneideri spent the entire day cycle inside the shelter. As toads did not engage in nocturnal activity in those areas with low thermal quality, the overall effectiveness of thermoregulation was, indeed, elevated. In conclusion, we showed that daily and seasonal variation in Tb of an anuran species is highly associated with their respective pattern of activity and may involve important physiological and ecological compromises.