Seasonal variation in thermal energetics of the Australian owlet-nightjar (Aegotheles cristatus)

Many birds living in regions with seasonal fluctuations in ambient temperatures (T_a) typically respond to cold by increasing insulation and adjusting metabolic rate. Seasonal variation in thermal physiology has not been studied for the Caprimulgiformes, an order of birds that generally have basal metabolic rates (BMR) lower than predicted for their body mass. We measured the metabolic rate and thermal conductance of Australian owlet-nightjars (Aegotheles cristatus) during summer and winter using open-flow respirometry. Within the thermoneutral zone (TNZ; 31.3 to 34.8 °C), there was no seasonal difference in BMR or thermal conductance (C), but body temperature was higher in summer- (38.2 ± 0.3 °C) than winter-acclimatized (37.1 ± 0.5 °C) birds. Below the TNZ, resting metabolic rate (RMR) increased linearly with decreasing T_a, and RMR and C were higher for summer- than winter-acclimatized birds. The mean mass-specific BMR of owlet-nightjars (1.27 mL O₂ g^− 1 h^− 1) was close to the allometrically predicted value for a 45 g Caprimulgiformes, but well below that predicted for birds overall. These results suggest that owlet-nightjars increase plumage insulation to cope with low winter T_a, which is reflected in the seasonal difference in RMR and C below the TNZ, rather than adjusting BMR.     

Experimental tests of a seasonally changing visual preference for habitat in a long-distance migratory shorebird

Migratory shorebirds show highly organized seasonal cycles in physiological and morphological traits (body mass and composition, plumage, hormone levels, etc.), which in captivity is accompanied by restless behaviour at times when free-living birds would start migration. We introduce the idea that seasonally changing preference for habitat could motivate migrants to embark on migration and that this cognitive process could also guide them to seasonally appropriate places. We explored this by testing whether red knots (Calidris canutus), which also in captivity maintain marked circannual phenotypic rhythms, show evidence of seasonal change in preference for pictures of seasonally appropriate habitats. We first developed a method to verify whether red knots are able to memorize and discriminate contrasting pictures projected by LCD projectors. This was followed by two different experiments in which we tested for a seasonally changing preference for breeding or non-breeding habitat. When carried out during the pre-breeding season, the red knots are expected to prefer pictures of mudflats, their non-breeding habitat. At the start of the breeding season, they should prefer pictures of the tundra breeding habitat. We established that knots are able to distinguish and memorize projected images. We failed to demonstrate the predicted change in vision-based habitat preference, but for reasons of test design we do not interpret this as a strong rejection of the hypothesis. Instead, we suggest that experiments with greater numbers of individuals tested once, perhaps in combination with the provision of additional cues such as smells and sounds, will help the development of these ideas further.     

Seasonal variations in basal metabolic rate, lower critical temperature and responses to temporary starvation in the arctic fox (Alopex lagopus) from Svalbard

Metabolic rates of four resting, post-absorptive male adult summer- and winter-adapted captive arctic foxes (Alopex lagopus) were recorded. Basal metabolic rates (BMR) varied seasonally with a 36% increase from winter to summer, while body mass was reduced by 17% in the same period. The lower critical temperature (T _1c) of the winter-adapted arctic fox was estimated to −7°C, whereas T _lc during summer was 5°C. The similarity of these values, which are much higher than hitherto assumed (e.g. Scholander et al. 1950b), is mainly due to a significantly (P<0.05) lower BMR in winter than in summer. Body core (stomach) temperature was stable, even at ambient temperatures as low as −45°C, but showed a significant (P<0.05) seasonal variation, being lower in winter (39.3±0.33°C) than in summer (39.8±0.16°C). The thermal conductivity of arctic fox fur was the same during both seasons, whereas the thermal conductance in winter was lower than in summer. This was reflected in an increase in fur thickness of 140% from summer to winter, and in a reduced metabolic response to ambient temperatures below T _lc in winter. Another four arctic foxes were exposed to three periods of forced starvation, each lasting 8 days during winter, when body mass is in decline. No significant reduction in mass specific BMR was observed during the exposure to starvation, and respiratory quotient was unchanged at 0.73±0.02 during the first 5 days, but dropped significantly (P<0.05) to 0.69±0.03 at day 7. Locomotor activity and body core (intraperitoneal) temperature was unaltered throughout the starvation period, but body mass was reduced by 18.5±2.1% during these periods. Upon re-feeding, locomotor activity was significantly (P<0.05) reduced for about 6 days. Energy intake was almost doubled, but stabilised at normal levels after 11 days. Body mass increased, but not to the level before the starvation episodes. Instead, body mass increased until it reached the reduced body mass of ad libitum fed control animals. This indicates that body mass in the arctic fox is regulated according to a seasonally changing set point.     

Moult and basal metabolic costs in males of two subspecies of stonechats: the European Saxicola torquata rubicula and the East African S. t. axillaris

The circannual patterns in resting metabolic rate (RMR) of males of two subspecies of stonechats, the European Saxicola torquata rubicula and the East African S. t. axillaris, are compared. As the birds from the two subspecies were raised and kept under comparable laboratory conditions, differences in metabolic rate between the two subspecies had to be genetically determined. RMR peaked during moult in both subspecies. During the rest of the year RMR was fairly constant in both subspecies and assumed to reflect basal metabolic rate (BMR). African stonechats had a 22% lower mass specific BMR than European stonechats, which is thought to reflect a genetical physiological adaptation to the differences in environmental circumstances they experience in the field. A low BMR makes an animal more susceptible to cold. Hence, the relatively high plumage mass in the African compared to the European stonechat may be functionally linked to its relatively low BMR. Moult costs, calculated from the plumage masses and the differences in RMR inside and outside the moult period, tended to be higher in the European compared to the African stonechats. These data and an interspecific comparison of moult costs over various species of birds support the earlier notion by Lindström et al. (1993) that moult costs are more closely linked with BMR than with body mass or rate of moult. The relation between moult costs and BMR and the fact that the efficiency of moult is extremely low (3.8 and 6.4% for European and African stonechats, respectively) suggest that the maintenance of specific tissues necessary for moult is a large cost factor. Alternatively, impaired insulation during moult may necessitate an increased metabolic capacity which may be associated with an increased RMR.     

Seasonal trends in body mass, food intake and resting metabolic rate, and induction of metabolic depression in arctic foxes (Alopex lagopus) at Svalbard

 Post-absorptive resting metabolic rates (RMRs), body mass and ad libitum food intake were recorded on an annual cycle in captive arctic foxes (Alopex lagopus) at Svalbard. During the light season in May and in the dark period in November, RMR during starvation and subsequent re-feeding were also measured. In contrast to earlier findings, the present study indicated a seasonal trend in post-absorptive RMR (in W · kg⁻¹ and W · kg^−0.75). The values in the light summer were 15% and 11% higher than the values in the dark winter, suggesting a physiological adaptation aiding energy conservation during winter in arctic foxes. Body mass and ad libitum food intake varied inversely through the year. A significant reduction in RMR (in W and W · kg^−0.75) with starvation (metabolic depression) was recorded both in May and November, indicating an adaptation to starvation in arctic foxes. The lack of metabolic depression during a period of starvation that was concomitant with extremely cold ambient temperatures in November 1994 indicates that metabolic responses to starvation may be masked by thermoregulatory needs. At very low ambient temperatures, arctic foxes may require increased heat production which cannot be achieved via below-average rates of metabolism. Accepted: 7 June 1999     

Temperature regulation in a burrow-nesting tropical seabird,the Wedge-tailed Shearwater (Puffinus pacificus)

Summary At low air temperatures (2.3–13.9°C), Wedge-tailed Shearwaters (Puffinus pacificus) shivered and their oxygen consumption increased to as much as 283% of the mean value (0.77 ml O₂/g·h) within the thermoneutral zone of air temperature (23–34°C). The minimal thermal conductance of the tissues and plumage was similar to the value predicted from the body mass (320.5 g). The oxygen consumption of the birds within their thermoneutral zone was lower than predictions based on body mass. At elevated air temperatures, the shearwaters panted at respiratory frequencies as high as 260 respirations/min; maximal respiratory frequencies were not invoked until the birds had become hyperthermic. During exposure to a hot environment, the oxygen consumption of the birds increased and in most instances the shearwaters were not able to lose heat equivalent to their concurrent metabolic heat production.Symbols and abbreviations BMR basal metabolic rate - C _total total thermal conductance - f respiratory frequency - TEWL total evaporative water loss - T _st stomach temperature - T _re rectal temperature     

Warm-up rates during arousal from torpor in heterothermic mammals: physiological correlates and a comparison with heterothermic insects

Summary This study examines the relationship between warm-up rate, body mass, metabolic rate, thermal conductance and normothermic body temperature in heterothermic mammals during arousal from torpor. Predictions based on the assumption that the energetic cost of arousal has been minimised are tested using data for 35 species. The observation that across-species warm-up rate correlates negatively with body mass is confirmed using a comparative technique which removes confounding effects due to the non-independence of species data due to shared common ancestry. Mean warm-up rate during arousal correlates negatively with basal metabolic rate and positively with the temperature difference through which the animal warms, having controlled for other factors. These results suggest that selection has operated to minimise the overall energetic, cost of warm-up. In contrast, peak warm-up rate during arousal correlates positively with peak metabolic rate during arousal, and negatively with thermal conductance, when body mass has been taken into account. These results suggest that peak warm-up rate is more sensitive to the fundamental processes of heat generation and loss. Although heterothermic marsupials have lower normothermic body temperatures and basal metabolic rates, marsupials and heterothermic eutherian mammals do not differ systematically in warm-up rate. Pre-flight warm-up rates in one group of endothermic insects, the bees, are significantly higher than predictions based on rates of arousal of a mammal of the same body mass.Abbreviations BMR basal metabolic rate - ICM independent comparisons method - MWR mean warm-up rate - PMR peak metabolic rate - PWR peak·warm-up rate - Tb_activity body temperature during activity - Tb_torpor body temperature during torpor - T _arousal increase in body temperature during arousal     

Life history varies with migratory distance in western sandpipers Calidris mauri

The propensity of migratory waders to remain on the non-breeding grounds during the arctic breeding season ("oversummer") in their first biological year of life ("juveniles") may be latitude, and thus migratory distance dependent. We compared the extent of preparation for northward migration of western sandpipers Calidris mauri spending the non-breeding season in México and Panamá during 1995–1998. During winter residency and premigratory periods, we measured body mass and scored the extent of dull basic versus bright alternate breeding plumage of captured juveniles and adults (second biological year or older), and obtained additional plumage scores from observations of uniquely colour banded birds. Nearly all western sandpipers in México prepared for northward migration by increasing body mass and moulting into breeding plumage. In Panamá, most adults prepared for migration, but few, if any, juveniles did so. Patterns of body mass and breeding plumage development do not generally support the hypothesis that oversummering by juveniles results directly from less efficient foraging or from resource competition with adults. We suggest instead that costs directly associated with migratory distance per se influence the life history strategies of sandpipers spending the non-breeding seasons at different latitudes. This latitudinal difference should interact with the well documented sex-ratio cline in non-breeding distribution (male western sandpipers predominating in northern parts of the range and females in southern parts). This suggests that females have more conservative life histories, prioritizing first year survivorship, relative to males that instead weight first-year breeding opportunities.     

Shorebirds' seasonal adjustments in thermogenic capacity are reflected by changes in body mass: how preprogrammed and instantaneous acclimation work together

Phenotypic flexibility in shorebirds has been studied mainly in the context of adjustments to migration and to quality of food; little is known on how birds adjust their phenotype to harsh winter conditions. We showed earlier that red knot (Calidris canutus islandica) can acclimate to cold by elevating body mass. This goes together with larger pectoral muscles, i.e., greater shivering machinery, and thus, better thermogenic capacity. Here, we present results of a yearlong experiment with indoor captive knots to determine whether this strategy is part of their natural seasonal phenotypic cycle. We maintained birds under three thermal regimes: constant cold (5 °C), constant thermoneutrality (25 °C) and natural seasonal variation between these extremes (9-22 °C). Each month we measured variables related to the birds' endurance to cold and physiological maintenance [body mass, thickness of pectoral muscles, summit metabolic rate (M(sum)), food intake, gizzard size, basal metabolic rate (BMR)]. Birds from all treatments expressed synchronized and comparable variation in body mass in spite of thermal treatments, with a 17-18% increase between the warmest and coldest months of the year; which appeared regulated by an endogenous driver. In addition, birds living in the cold exhibited a 10% higher average body mass than did those maintained at thermoneutrality. Thickness of the pectoral muscle tracked changes in body mass in all treatments and likely contributed to greater capacity for shivering in heavier birds. Consequently, M(sum) was 13% higher in cold-acclimated birds compared to those experiencing no thermoregulation costs. However, our data also suggest that part of maximal heat production comes from nonshivering processes. Birds facing cold conditions ate up to 25% more food than did birds under thermoneutral conditions, yet did not develop larger gizzards. Seasonal variation in BMR followed changes in body mass, probably reflecting changes in mass of metabolically active tissues. Just as cold-exposed birds, red knots in the variable treatment increased body mass in winter, thereby improving cold endurance. During summer, however, they maintained a lower body mass and thermogenic capacity compared to cold-exposed birds, similar to individuals kept at thermoneutrality. We conclude that red knots acclimate to seasonal variations in ambient temperature by modulating body mass, combining a preprogrammed increase in mass during winter with a capacity for fine-tuning body mass and thermogenic capacity to temperature variations.     

Effects of long-term captivity on thermoregulation,metabolism and ventilation of the southern brown bandicoot (Marsupialia: Peramelidae)

Thermoneutral metabolic and ventilatory parameters were measured every 3 months over 2 years for southern brown bandicoots held in captivity, and from a nearby reserve. Captive bandicoots were 130 g (9.9%) heavier than wild bandicoots. Long-term captivity had no effect on body temperature, basal metabolic rate (oxygen consumption), thermal conductance or respiratory ventilation, but there was an effect on carbon dioxide production, respiratory exchange ratio and total evaporative water loss (values were between 15 and 25% higher for captive than for wild bandicoots). Diet may be influencing these aspects of captive bandicoot physiology; the diet of captive bandicoots would be considerably different to that of wild bandicoots. Water availability seems to have a minimal effect. This study has important implications regarding physiological measurement for captive and wild mammals. For bandicoots at least, captive animals are equivalent to wild animals for some physiological parameters at thermoneutrality (body temperature, resting metabolic rate and thermal conductance), but not others.     

Daily rhythm of oxygen consumption and thermoregulatory responses in some European winter- or summer-acclimatized finches at different ambient temperatures

The oxygen consumption of European finches, the siskin (Carduelis spinus), the brambling (Fringilla montifringilla), the bullfinch (Pyrhulla pyrhulla), the greenfinch (Carduelis chloris) and the hawfinch (Coccothraustes coccothraustes), was recorded continuously while ambient temperature was decreased stepwise from +30 down to-75°C. The oxygen consumption, body temperature (telemetrically), and shivering (integrated pectoral electromyography) of greenfinches were measured simultaneously at ambient temperatures between +30 and-75°C. Maximum heat production, cold limit, lower critical temperature, basal metabolic rate and thermal conductance (of the greenfinch) were determined. The diurnal variation of oxygen consumption of siskins and greenfinches was recorded at thermoneutrality and below the thermoneutral zone in winter- and summer-acclimatized birds. The diurnal variation of body temperature and thermal conductance of greenfinches were also determined. The diurnal variation of heat production was not seasonal or temperature dependent in the siskin and in the greenfinch. Nocturnal reduction of oxygen consumption saved 15–33% energy in the siskin and greenfinch. Body temperature of the greenfinch was lowered by 2.5–3.4°C. The nocturnal reduction of thermal conductance in the greenfinch was 39–48%. The basal metabolic rate was lowest in the largest bird (hawfinch) and highest in the smallest bird (siskin). The values were in the expected range. The heat production capacity of finches in winter was 4.7 times basal metabolic rate in the siskin, 4.2 times in the brambling, 3.5 times in the greenfinch and 2.9 times in the bullfinch and hawfinch. The heat production capacity of the siskin and greenfinch was not significantly lower in summer. The cold limit temperatures (°C) in winter were-61.2 in the siskin,-41.3 in the greenfinch,-37.0 in the bullfinch,-35.7 in the brambling and-28.9 in the hawfinch. The cold limit was 14.3°C higher in summer than in winter in the siskin and 8.7°C in the greenfinch. Thermal insulation of the greenfinch was significantly better in winter than in summer. The shivering of the greenfinch increased linearly when ambient temperature was decreased down to-40°C. Maintenance of shivering was coincident with season. In severe cold integrated pectoral electromyography did not correlate with oxygen consumption as expected. The possible existence of non-shivering thermogenesis in birds is discussed. It is concluded that the acclimatization of European finches is primarily metabolic and only secondly affected by insulation.Abbreviations AAT avian adipose tissue - bm body mass - BMR basal metabolic rate - C _t thermal conductance - EMG electromyogram - HP heat production - HP _max maximum heat production - MR metabolic rate - NST non-shivering thermogenesis - RMR resting metabolic rate - RQ respiratory quotient - T _a ambient temperature - T _b body temperature - T _c colonic temperature - T _1c lower critical temperature - TNZ thermoneutral zone - T _st shivering threshold temperature - V oxygen consumption     

Summer acclimatization in the short-tailed field vole,Microtus agrestis

We investigated the changes that occurred in basal and noradrenaline-induced metabolic rate, body temperature and body mass in short-tailed field voles,Microtus agrestis, during exposure to naturally increasing photoperiod and ambient temperature. These parameters were first measured in winter-acclimatized voles (n=8) and then in the same voles which had been allowed to seasonally acclimatize to photoperiod and ambient temperature (6 months later). Noradrenaline induced metabolic rate, basal metabolic rate and nonshivering thermogenesis were significantly higher in winter-acclimatized compared to summer-acclimatized voles. There was a significant positive relationship between basal metabolic rate and noradrenaline-induced metabolic rate. Body mass was significantly higher in summer-acclimatized compared to winter-acclimatized voles. There was a significant positive relationship between body mass and noradrenaline-induced metabolic rate in both winter-acclimalized and summer-acclimatized voles; however, there was no relationship between basal metabolic rate and body mass in either seasonal group of voles. Body temperature after measurements of basal metabolic rate was not significantly different in the seasonal cohorts of voles. However, body temperature was significantly higher in winter-acclimatized compared to summer-acclimatized voles after injection of noradrenaline. Previously we have found that a long photoperiod was not a sufficient stimulus to reduce thermogenic capacity in winter-acclimatized voles during cold exposure, since basal metabolic rate increased to compensate for a reduction in regulatory nonshivering thermogenesis. Here we found that a combination of increased ambient temperature and photoperiod did significantly reduce thermogenic capacity in winter-acclimatized voles. This provided evidence that the two aspects of non-shivering thermogenesis, obligatory and regulatory, are stimulated by different exogenous cues. Summer acclimatization in the shorttailed field vole is manifest as a significant decrease in both basal and noradrenaline-induced metabolic rate, combined with a significant increase in body mass.Abbreviations ANCOV A analysis of covariance - BAT brown adipose tissue - BM body mass - BMR basal metabolic rate - NST non-shivering thermogenesis - NA noradrenaline - V the maximum V recorded following mass specific injection of noradrenaline - V the maximum V recorded following mass specific injection of saline - T _a ambient temperature - T _b rectal body temperature - T _1c lower critical temperature - UCP uncoupling protein - V oxygen consumption     

Ambient temperature does not affect fuelling rate in absence of digestive constraints in long-distance migrant shorebird fuelling up in captivity

Pre-flight fuelling rates in free-living red knots Calidris canutus, a specialized long-distance migrating shorebird species, are positively correlated with latitude and negatively with temperature. The single published hypothesis to explain these relationships is the heat load hypothesis that states that in warm climates red knots may overheat during fuelling. To limit endogenous heat production (measurable as basal metabolic rate BMR), birds would minimize the growth of digestive organs at a time they need. This hypothesis makes the implicit assumption that BMR is mainly driven by digestive organ size variation during pre-flight fuelling. To test the validity of this assumption, we fed captive knots with trout pellet food, a diet previously shown to quickly lead to atrophied digestive organs, during a fuelling episode. Birds were exposed to two thermal treatments (6 and 24°C) previously shown to generate different fuelling rates in knots. We made two predictions. First, easily digested trout pellet food rather than hard-shelled prey removes the heat contribution of the gut and would therefore eliminate an ambient temperature effect on fuelling rate. Second, if digestive organs were the main contributors to variations in BMR but did not change in size during fuelling, we would expect no or little change in BMR in birds fed ad libitum with trout pellets. We show that cold-acclimated birds maintained higher body mass and food intake (8 and 51%) than warm-acclimated birds. Air temperature had no effect on fuelling rate, timing of fuelling, timing of peak body mass or BMR. During fuelling, average body mass increased by 32% while average BMR increased by 15% at peak of mass and 26% by the end of the experiment. Our results show that the small digestive organs characteristic of a trout pellet diet did not prevent BMR from increasing during premigratory fuelling. Our results are not consistent with the heat load hypothesis as currently formulated.     

Body temperature and metabolic rate during natural hypothermia in endotherms

During daily torpor and hibernation metabolic rate is reduced to a fraction of the euthermic metabolic rate. This reduction is commonly explained by temperature effects on biochemical reactions, as described by Q ₁₀ effects or Arrhenius plots. This study shows that the degree of metabolic suppression during hypothermia can alternatively be explained by active downregulation of metabolic rate and thermoregulatory control of heat production. Heat regulation is fully adequate to predict changes in metabolic rate, and Q ₁₀ effects are not required to explain the reduction of energy requirements during hibernation and torpor.Abbreviations BMR basal metabolic rate - BW body weight - C thermal conductance - C_HL thermal conductance as derived from HL - C_HP thermal conductance as derived from HP - HL heat loss - HP heat production - MR metabolic rate - RQ respiratory quotient - T_a ambient temperature - T_b body temperature     

Gizzard and other lean mass components increase, yet Basal Metabolic Rates decrease, when red knots Calidris canutus are shifted from soft to hard-shelled food

We measured basal metabolic rate (BMR), body mass, lean mass, and gizzard mass of captive red knots Calidris canutus islandica maintained on a trout chow diet (soft-texture, low ash and water content) for several years and then shifted to small mussels Mytilus edulis (hard-texture, high ash and water content). During a 3-week period of feeding on mussels, body mass, lean mass, and gizzard mass increased 7.3 g (+7%), 10.5 g (+12%), and 4.9 g (+213%), respectively, yet BMR decreased from 0.96 to 0.89 W (−8%). Under the new mussel regime, red knots must have reduced the metabolic intensity of some of the tissues. This suggests that the experimental red knots experienced the transition to a mussel diet as stressful and energy limiting, resulting in an energy-saving strategy by reducing BMR in spite of hypertrophy of the gizzard and other organs.     

Travelling on a budget: predictions and ecological evidence for bottlenecks in the annual cycle of long-distance migrants

Long-distance migration, and the study of the migrants who undertake these journeys, has fascinated generations of biologists. However, many aspects of the annual cycles of these migrants remain a mystery as do many of the driving forces behind the evolution and maintenance of the migrations themselves. In this article we discuss nutritional, energetic, temporal and disease-risk bottlenecks in the annual cycle of long-distance migrants, taking a sandpiper, the red knot Calidris canutus, as a focal species. Red knots have six recognized subspecies each with different migratory routes, well-known patterns of connectivity and contrasting annual cycles. The diversity of red knot annual cycles allows us to discuss the existence and the effects of bottlenecks in a comparative framework. We examine the evidence for bottlenecks focusing on the quality of breeding plumage and the timing of moult as indicators in the six subspecies. In terms of breeding plumage coloration, quality and timing of prealternate body moult (from non-breeding into breeding plumage), the longest migrating knot subspecies, Calidris canutus rogersi and Calidris canutus rufa, show the greatest impact of bottlenecking. The same is true in terms of prebasic body moult (from breeding into non-breeding plumage) which in case of both C. c. rogersi and C. c. rufa overlaps with southward migration and may even commence in the breeding grounds. To close our discussion of bottlenecks in long-distance migrants, we make predictions about how migrants might be impacted via physiological 'trade-offs' throughout the annual cycle, using investment in immune function as an example. We also predict how bottlenecks may affect the distribution of mortality throughout the annual cycle. We hope that this framework will be applicable to other species and types of migrants, thus expanding the comparative database for the future evaluation of seasonal selection pressures and the evolution of annual cycles in long-distance migrants. Furthermore, we hope that this synthesis of recent advancements in the knowledge of red knot annual cycles will prove useful in the ongoing attempts to model annual cycles in migratory birds.     

Inter-annual consistency in the fluctuating energy requirements of captive harp seals Phoca groenlandica

To examine seasonal and inter-annual shifts in the energy requirements of captive harp seals (Phoca groenlandica), metabolic rate and body temperature were repeatedly measured over the annual cycle. Seasonal shifts were evident in both parameters, with spring and summer (April to September) peaks decreasing throughout fall and winter (October to March). Seasonal changes in oxygen consumption concur with earlier published reports and data presented here validate these trends inter-annually. The standard metabolic rates (SMRs) of all seals were lower than predicted during fall/winter, but were indistinguishable from expected rates during spring/summer. Although individual variation in metabolic rate was largely independent of changes in both total body mass and predicted total body fat over the year, such variation was more closely related to changes in predicted lean body mass. Both deep rectal and core body temperatures co-varied with metabolic rate, perhaps indicating a metabolic defense of fat for thermoregulation. The implications of these shifting basal requirements are considered in the light of calculating the impact of the harp seal herd on fisheries resources in the Northwest Atlantic. Received: 18 December 1996 / Accepted: 30 April 1997     

Seasonal changes in the thermoenergetics of the marsupial sugar glider, Petaurus breviceps

Little information is available on seasonal changes in thermal physiology and energy expenditure in marsupials. To provide new information on the subject, we quantified how body mass, body composition, metabolic rate, maximum heat production, body temperature and thermal conductance change with season in sugar gliders (Petaurus breviceps) held in outdoor aviaries. Sugar gliders increased body mass in autumn to a peak in May/June, which was caused to a large extent by an increase in body fat content. Body mass then declined to minimum values in August/September. Resting metabolic rate both below and above the thermoneutral zone (TNZ) was higher in summer than in winter and the lower critical temperature of the TNZ occurred at a higher ambient temperature (Ta) in summer. The basal metabolic rate was as much as 45% below that predicted from allometric equations for placental mammals and was about 15% lower in winter than in summer. In contrast, maximum heat production was raised significantly by about 20% in winter. This, together with an approximately 20% decrease in thermal conductance, resulted in a 13 degrees C reduction of the minimum effective Ta gliders were able to withstand. Our study provides the first evidence that, despite the apparent lack of functional brown adipose tissue, sugar gliders are able to significantly increase heat production in winter. Moreover, the lower thermoregulatory heat production at most TaS in winter, when food in the wild is scarce, should allow them to reduce energy expenditure.     

Male territorial aggression and androgen modulation in high latitude populations of the Sooty, Passerella iliaca sinuosa, and Red Fox Sparrow, Passerella iliaca zaboria

The Fox Sparrows, Passerella iliaca, include multiple groups and subspecies distributed at several latitudes from the Alaskan arctic to the southwestern United States. As such, this species represents a potential model for investigating latitudinal variation in androgen secretion and aggressive territoriality in male passerines. Breeding male Fox Sparrows from two subspecies within two groups, the Sooty Fox Sparrow, P. i. sinuosa, and the Red Fox Sparrow, P. i. zaboria, were assessed for aggressive territoriality and androgen responsiveness at multiple latitudes in arctic and subarctic Alaska. Subarctic Sooty Fox Sparrows had higher circulating androgen levels in the early (8.54 ng/ml) versus mid–late breeding season (2.44 ng/ml). Males in the mid–late breeding season did not up-regulate androgen secretion in response to social challenge, but were aggressive and spent more time within 5 m of a decoy during a simulated territorial intrusion (STI) than early breeding males. Male subarctic Red Fox Sparrows had slightly higher circulating androgen levels (2.29 ng/ml) than arctic males (1.10 ng/ml) in the mid–late breeding season. However, androgen levels were not correlated with blood collection time after a social challenge in either group, suggesting that neither arctic nor subarctic males up-regulate androgen secretion during the mid–late breeding period. Arctic males spent more time within 5 m of a decoy and sang less than subarctic males during an STI in the mid–late breeding season. These findings demonstrate that the Fox Sparrow is a tractable model for investigating the latitudinal regulation of aggressive territoriality and androgen responsiveness in passerines.     

Contrasting extreme long‐distance migration patterns in bar‐tailed godwits Limosa lapponica

Migrating birds make the longest non‐stop endurance flights in the animal kingdom. Satellite technology is now providing direct evidence on the lengths and durations of these flights and associated staging episodes for individual birds. Using this technology, we compared the migration performance of two subspecies of bar‐tailed godwit Limosa lapponica travelling between non‐breeding grounds in New Zealand (subspecies baueri) and northwest Australia (subspecies menzbieri) and breeding grounds in Alaska and eastern Russia, respectively. Individuals of both subspecies made long, usually non‐stop, flights from non‐breeding grounds to coastal staging grounds in the Yellow Sea region of East Asia (average 10 060 ± SD 290 km for baueri and 5860 ± 240 km for menzbieri). After an average stay of 41.2 ± 4.8 d, baueri flew over the North Pacific Ocean before heading northeast to the Alaskan breeding grounds (6770 ± 800 km). Menzbieri staged for 38.4 ± 2.5 d, and flew over land and sea northeast to high arctic Russia (4170 ± 370 km). The post‐breeding journey for baueri involved several weeks of staging in southwest Alaska followed by non‐stop flights across the Pacific Ocean to New Zealand (11 690 km in a complete track) or stopovers on islands in the southwestern Pacific en route to New Zealand and eastern Australia. By contrast, menzbieri returned to Australia via stopovers in the New Siberian Islands, Russia, and back at the Yellow Sea; birds travelled on average 4510 ± 360 km from Russia to the Yellow Sea, staged there for 40.8 ± 5.6 d, and then flew another 5680–7180 km to Australia (10 820 ± 300 km in total). Overall, the entire migration of the single baueri godwit with a fully completed return track totalled 29 280 km and involved 20 d of major migratory flight over a round‐trip journey of 174 d. The entire migrations of menzbieri averaged 21 940 ± 570 km, including 14 d of major migratory flights out of 154 d total. Godwits of both populations exhibit extreme flight performance, and baueri makes the longest (southbound) and second‐longest (northbound) non‐stop migratory flights documented for any bird. Both subspecies essentially make single stops when moving between non‐breeding and breeding sites in opposite hemispheres. This reinforces the critical importance of the intertidal habitats used by fuelling godwits in Australasia, the Yellow Sea, and Alaska.