Mass–size relationships,starvation and recovery in an engorging feeder

The relationships of larval nutritional resources with adult body size, starvation resistance and reproductive decisions are not always clear. Burying beetle larvae with inadequate nutrition are hypothesized to develop into relatively large adults that are able to contest for breeding resources. The trade‐off is that the emerging adult has minimal energy reserves and is more susceptible to starvation, and must gain proportionately more weight after emerging. These hypotheses are investigated in Nicrophorus orbicollis Say. In addition, sex differences in size–mass ratios as well as starvation and recovery in reproductively mature females are examined. The larval mass to adult size ratio is similar in male and female N. Orbicollis and, contrary to prediction, small larvae do not result in adults that are relatively large in size for their mass. Emerging adults of lesser mass resist starvation less well, as expected. Emerging adults of smaller pronotal size gain more relative mass but less absolute mass than larger adults. In reproductively mature adult females, recovery from food deprivation is rapid, with most if not all the weight that is lost during a 9–10‐day starvation period being re‐gained within 1 day of engorging. The ability to gain weight rapidly and regulate body mass provides a nutritional framework for understanding the larva to adult transition and the reproductive and parenting decisions of burying beetles that otherwise would appear to be of too high risk.     

Risk‐sensitive reproductive allocation: fitness consequences of body mass losses in two contrasting environments

For long‐lived organisms, the fitness value of survival is greater than that of current reproduction. Asymmetric fitness rewards suggest that organisms inhabiting unpredictable environments should adopt a risk‐sensitive life history, predicting that it is adaptive to allocate resources to increase their own body reserves at the expense of reproduction. We tested this using data from reindeer populations inhabiting contrasting environments and using winter body mass development as a proxy for the combined effect of winter severity and density dependence. Individuals in good and harsh environments responded similarly: Females who lost large amounts of winter body mass gained more body mass the coming summer compared with females losing less mass during winter. Additionally, females experienced a cost of reproduction: On average, barren females gained more body mass than lactating females. Winter body mass development positively affected both the females' reproductive success and offspring body mass. Finally, we discuss the relevance of our findings with respect to scenarios for future climate change.     

Does avian winter fat storage integrate temperature and resource conditions? A long‐term study

Theoretical models of short‐term avian behaviour suggest that small birds adaptively balance the ecological costs and benefits of winter fat to maximize survival probability. When low starvation risk eliminates benefit but not cost of fat, birds are leaner than when under high starvation risk. Most models focus on single factors affecting starvation risk and subsequent choice of adaptive body mass; however, in complex environments, more than one factor affects starvation risk. To test for multiple interacting factors affecting fat reserves, long‐term geographical data on winter fat in a ground‐feeding finch, the dark‐eyed junco Junco hyemalis were analyzed. Two measures of fat were used: (1) visible subcutaneous fat class, and (2) body mass residuals left after age, sex and wing length effects were factored out. Site means for fat measures were obtained from juncos visiting supplemental feeding sites in midwest and northwest North America. In backward elimination regression of fat class, the temperature‐snowfall interaction term and its constituent variables, proximate temperature (averaged over capture day and the preceding ten days) and snowfall (frequency over the same time interval) were significant explanators of variation. Snowfall frequency is considered to be a surrogate measure of resource deterioration. The interaction term, also found in backward regression of body mass residuals, showed that as temperature declined at low snowfall frequency, less fat was deposited than when temperature declined at high snowfall frequency. Thus, in a recently cold environment suggesting relatively high resource predictability, perceived starvation risk is low, and less costly fat is needed to reduce starvation risk compared to a cold and unpredictable resource environment. The analysis of mass residuals also yielded a significant effect of daylength, suggesting an underlying fattening programme independent of proximate environmental conditions. A longitudinal study of junco fat stores indicated that individual environmental responses contributed significantly to midwinter fat peaks. These results agree with predictions of a synergistic model of adaptive fat regulation in small birds by suggesting that a ground‐feeding bird may maximize winter survival probability by integrating multiple environmental factors affecting starvation risk.     

Changes in body condition of hibernating bats support the thrifty female hypothesis and predict consequences for populations with white-nose syndrome

White-nose syndrome (WNS) is a new disease of bats that has devastated populations in eastern North America. Infection with the fungus, Geomyces destructans, is thought to increase the time bats spend out of torpor during hibernation, leading to starvation. Little is known about hibernation in healthy, free-ranging bats and more data are needed to help predict consequences of WNS. Trade-offs presumably exist between the energetic benefits and physiological/ecological costs of torpor, leading to the prediction that the relative importance of spring energy reserves should affect an individual''s use of torpor and depletion of energy reserves during winter. Myotis lucifugus mate during fall and winter but females do not become pregnant until after spring emergence. Thus, female reproductive success depends on spring fat reserves while male reproductive success does not. Consequently, females should be “thrifty” in their use of fat compared to males. We measured body condition index (BCI; mass/forearm length) of 432 M. lucifugus in Manitoba, Canada during the winter of 2009/2010. Bats were captured during the fall mating period (n = 200), early hibernation (n = 125), and late hibernation (n = 128). Adult females entered hibernation with greater fat reserves and consumed those reserves more slowly than adult males and young of the year. Consequently, adult females may be more likely than males or young of the year to survive the disruption of energy balance associated with WNS, although surviving females may not have sufficient reserves to support reproduction.     

Nocturnal loss of body reserves reveals high survival risk for subordinate great tits wintering at extremely low ambient temperatures

Winter acclimatization in birds is a complex of several strategies based on metabolic adjustment accompanied by long-term management of resources such as fattening. However, wintering birds often maintain fat reserves below their physiological capacity, suggesting a cost involved with excessive levels of reserves. We studied body reserves of roosting great tits in relation to their dominance status under two contrasting temperature regimes to see whether individuals are capable of optimizing their survival strategies under extreme environmental conditions. We predicted less pronounced loss of body mass and body condition and lower rates of overnight mortality in dominant great tits at both mild and extremely low ambient temperatures, when ambient temperature dropped down to ?43 °C. The results showed that dominant great tits consistently maintained lower reserve levels than subordinates regardless of ambient temperature. However, dominants responded to the rising risk of starvation under low temperatures by increasing their body reserves, whereas subdominant birds decreased reserve levels in harsh conditions. Yet, their losses of body mass and body reserves were always lower than in subordinate birds. None of the dominant great tits were found dead, while five young females and one adult female were found dead in nest boxes during cold spells when ambient temperatures dropped down to ?43 °C. The dead great tits lost up to 23.83 % of their evening body mass during cold nights while surviving individuals lost on average 12.78 % of their evening body mass. Our results show that fattening strategies of great tits reflect an adaptive role of winter fattening which is sensitive to changes in ambient temperatures and differs among individuals of different social ranks.     

Experimental evidence of cost of lactation in a low risk environment for a long-lived mammal

In a previous experiment we have documented that organisms adopt a risk-sensitive reproductive allocation when summer reproductive investment competes with survival in the coming winter ( Bårdsen et al. 2008 ). This tradeoff is present through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis experimentally on female reindeer experiencing stable and benign winter feeding conditions. Additional supplementary feeding and removal of newborns represented two sets of experimental manipulations. Females in the supplementary feeding group increased more in winter body mass relative to control females. This manipulation, however, did not have any effect on summer body mass development for neither females nor offspring, but we found a positive effect of feeding on offspring birth mass for smaller females. In contrast, offspring removal did have a positive effect on summer body mass development as females in this group were larger in the autumn relative to control females. In essence, we documented two immediate effects as: (1) supplementary feeding did have a positive effect on spring body mass for smaller females; and (2) offspring removal did increase the female summer somatic growth as this had a positive effect on female autumn body mass. Additionally, we tested for lagged effects, but we could not document any biologically significant effects of neither manipulation in the coming spring. The fact that we only found rather weak effects of both manipulations was as expected for risk sensitive individuals experiencing benign environmental conditions over many years.     

Loss of body mass under predation risk: cost of antipredatory behaviour or adaptive fit-for-escape?

Predation risk may compromise the ability of animals to acquire and maintain body reserves by hindering foraging efficiency and increasing physiological stress. Locomotor performance may depend on body mass, so losing mass under predation risk could be an adaptive response of prey to improve escape ability. We studied individual variation in antipredatory behaviour, feeding rate, body mass and escape performance in the lacertid lizard Psammodromus algirus. Individuals were experimentally exposed to different levels of food availability (limited or abundant) and predation risk, represented by reduced refuge availability and simulated predator attacks. Predation risk induced lizards to reduce conspicuousness behaviourally and to avoid feeding in the presence of predators. If food was abundant, alarmed lizards reduced feeding rate, losing mass. Lizards supplied with limited food fed at near-maximum rates independently of predation risk but lost more mass when alarmed; thus, mass losses experienced under predation risk were higher than those expected from feeding interruption alone. Although body mass of lizards varied between treatments, no component of escape performance measured during predator attacks (endurance, speed, escape strategy) was affected by treatments or by variations in body mass. Thus, the body mass changes were consistent with a trade-off between gaining resources and avoiding predators, mediated by hampered foraging efficiency and physiological stress. However, improved escape efficiency is not required to explain mass reduction upon predator encounters beyond that expected from feeding interruption or predation-related stress. Therefore, the idea that animals may regulate body reserves in relation to performance demands should be reconsidered.     

Sex, diet and feeding method-related differences in body condition in the Oystercatcher Haematopus ostralegus

Measurements were made of Oystercatcher corpses to find the most reliable measure of body condition in live birds. Total body weight was as accurate as muscle thickness in assessing the weight of the pectoral muscles, a major store of protein reserves. Total body weight was also a good indicator of fat reserves. Wing‐length was the most reliable measure of body size. Body condition in live birds was measured as an individual's residual weight derived from a general linear model of (loge) body weight against age, month, (loge) wing‐length and time since capture. On the Exe estuary in autumn, there was no significant difference in body condition between same‐age Oystercatchers of different diets and feeding methods. In the winter months, however, body condition differed significantly between mussel feeders using different feeding methods, and between mussel feeders and birds that fed on worms and clams. Differences between diet/feeding methods were not the same for each age group. Birds with the highest mean body condition index in adults were mussel hammerers, in immatures mussel stabbers and in juveniles, worm/clam feeders. No significant difference in body condition was found between the sexes in any age group in autumn. In winter, when birds with the same diet/feeding method were considered, females were significantly heavier than males. However, when birds of all diets and feeding methods were combined, weight differences between the sexes were not apparent. We examine reasons why birds with certain diets and feeding methods may be heavier than others in winter, and why females are sometimes heavier than males. We conclude that, in most cases, it is due to differences between birds in their ability to achieve a common target weight. However, we conclude that adult females try to achieve a higher target weight than males, probably as a premigratory and prebreeding strategy. In the light of this, we consider the consequences for adult females, in terms of reproductive success and survival, of specializing in diets and feeding methods associated with lower body weights in winter.     

Intermittent fasting during winter and spring affects body composition and reproduction of a migratory duck

We compared food intake, body mass and body composition of male and female black ducks (Anas rubripes) during winter (January-March). Birds were fed the same complete diet ad libitum on consecutive days each week without fasting (control; nine male; nine female) or with either short fasts (2 day x week(-1); nine male; nine female), or long fasts (4 day x week(-1); eleven male; twelve female). We continued treatments through spring (March-May) to measure the effect of intermittent fasts on body mass and egg production. Daily food intake of fasted birds was up to four times that of unfasted birds. Weekly food intake of males was similar among treatments (364 g x kg(-1) x week(-1)) but fasted females consumed more than unfasted females in January (363 g x kg(-1) x week(-1) vs. 225 g x kg(-1) x week(-1)). Although both sexes lost 10-14% body mass, fasted females lost less mass and lipid than unfasted females during winter. Total body nitrogen was conserved over winter in both sexes even though the heart and spleen lost mass while the reproductive tract and liver gained mass. Intermittent fasting increased liver, intestinal tissue and digesta mass of females but not of males. Fasting delayed egg production in spring but did not affect size, fertility or hatching of the clutch. Females on long fasts were still heavier than controls after laying eggs. Thus black ducks combine flexibility of food intake with plasticity of digestive tract, liver and adipose tissue when food supply is interrupted during winter. Females modulate body mass for survival and defer reproduction when food supply is interrupted in spring.     

Adaptations of the raccoon dog (Nyctereutes procyonoides) to wintering--effects of restricted feeding or periodic fasting on lipids, sex steroids and reproduction

The raccoon dog (Nyctereutes procyonoides) is an omnivorous canid with autumnal hyperphagia and fattening followed by mid-winter passivity and fasting in boreal latitudes with seasonal snow cover. The effects of two different feeding levels (400 or 200 kcal/animal/d) or fasting (5-week fasting+1-week feeding+3-week fasting) on plasma lipids, sex steroids and reproductive success of farm-bred raccoon dogs (n=60 females and 24 males) were studied in winter. The body masses, body mass indices (BMIs) and levels of plasma triacylglycerols (TG), total cholesterol and low- and high-density lipoprotein cholesterol did not differ between the fed and the restrictively fed animals. During fasting, the plasma TG concentrations increased and the BMIs decreased, indicating the release of fatty acids from adipose tissue. After the fasting periods, the levels of plasma cholesterol and high-density lipoprotein cholesterol increased, whereas the TG levels decreased indicating the rebuilding of energy reserves. The fact that the different wintertime feeding regimes had no impact on the plasma glucose, total protein, cortisol, estradiol, progesterone or testosterone levels, or on the reproductive success, indicates versatile adaptive capacity in the species.     

Quantifying energetic and fitness consequences of seasonal heterothermy in an Arctic ungulate

Animals have adapted behavioral and physiological strategies to conserve energy during periods of adverse conditions. Heterothermy is one such adaptation used by endotherms. While heterothermy—fluctuations in body temperature and metabolic rate—has been shown in large vertebrates, little is known of the costs and benefits of this strategy, both in terms of energy and in terms of fitness. Hence, our objective was to model the energetics of seasonal heterothermy in the largest Arctic ungulate, the muskox (Ovibos moschatus), using an individual‐based energy budget model of metabolic physiology. We found that the empirically based drop in body temperature (winter max ~−0.8°C) overwinter in adult females resulted in substantial fitness benefits in terms of reduced daily energy expenditure and body mass loss. Body mass and energy reserves were 8.98% and 14.46% greater in modeled heterotherms compared to normotherms by end of winter. Based on environmental simulations, we show that seasonal heterothermy can, to some extent, buffer the negative consequences of poor prewinter body condition or reduced winter food accessibility, leading to greater winter survival (+20%–30%) and spring energy reserves (+10%–30%), and thus increased probability of future reproductive success. These results indicate substantial adaptive short‐term benefits of seasonal heterothermy at the individual level, with potential implications for long‐term population dynamics in highly seasonal environments.     

Nutrition,sex and season contribute to variation in fat and glycerol levels in the long‐lived moth Caloptilia fraxinella

The ash leaf cone roller Caloptilia fraxinella Ely (Lepidoptera: Gracillariidae) is an invasive leaf‐mining moth pest of horticultural ash Fraxinus spp. in the Canadian Prairie Provinces. Caloptilia fraxinella overwinter as adults in reproductive diapause and mating occurs after overwintering in the spring. The effect of a carbohydrate food source on fat and glycerol reserves throughout the long adult life stage of this moth is investigated. Insects collected as pupae are given access to either water or sugar water upon adult eclosion. Moths held under the different feeding regimes are sampled before (summer and autumn) and after overwintering in the spring. Analysis of either glycerol or lipid content is conducted for male and female moths from each collection period. Both moth weight and glycerol concentration are affected by moth sex, food regime and season of collection. Although female moths are heavier than males, a higher glycerol concentration occurs in males. Moths fed sugar are heavier and have a higher glycerol concentration than water‐fed moths late in reproductive diapause and after overwintering. Moths collected in the spring after overwintering are lighter and have a lower glycerol content than moths collected before winter. There is a significant influence of feeding regime and season on moth body lipid content, with sugar‐fed moths having more fat than water‐fed moths; however, this difference is smaller in the summer than the autumn or spring. An initial understanding of the overwintering biology and diapause of this pest is provided in the present study.     

Energetic consequences of seasonal breeding in female Japanese macaques (Macaca fuscata)

Japanese macaques inhabit a relatively cold environment and females of this species could have developed strategies of energy economy to face the sometimes-harsh seasonal conditions of temperate climates, as well as reproductive costs, and thus regulate their energy balance. Here, we explore the relationship between nutritional status, body composition, seasonality, and reproductive status using isotope-labeled water, anthropometric measurements, and leptin assays from 14 captive female Japanese macaques. Our results indicated that body mass provided the best predictor of fat-free mass and fat mass. These females varied in estimated percent body fat between 8 and 25% (18% on average at the beginning of the mating season and 13% during the birth season). Higher body mass and body fat content were observed at the beginning of the mating season, which supports the hypothesis that individual females need to attain a sufficient physical condition to cover energy costs associated with mating activity, and to survive under severe ecological conditions in winter with high thermoregulatory costs. We found a relationship between conception rates and energetic condition or body fat, with females that conceived during one mating season being fatter after the end of their previous mating season. Together, these results suggest that, even in captive settings with constant food availability, seasonal breeding entails relatively high energy costs, and that females with higher energy status could invest more in reproductive activities and could afford to reproduce more rapidly.     

Effect of winter feeding and starvation on the growth performance of young‐of‐year (YOY) great sturgeon,Huso huso

The objective of this study was to evaluate the feeding rate of the great sturgeon (Huso huso) young of the year (YOY) and to investigate the effects of different feeding rates in maintaining the weight of fish during short periods of winter starvation. Six feeding rates of 0.2, 0.4, 0.6, 0.8, 1.0% body weight (BW) day^?1 and feeding to satiation were considered for the first experiment. Each feeding rate was randomly assigned to three replicate tanks, with continuous feeding throughout a 5‐week winter period of water temperatures below 10°C. Fifteen fish were held in each of 18 tanks with an average initial body weight of 219.6 ± 6.9 g. After 5 weeks of feeding, the best performance was observed in fish fed 1% BW day^?1, but negative growth was observed in fish fed 0.2% BW day^?1. In the second experiment, fish were deprived of feed for 3 weeks at winter temperatures. Weights and condition factors of all fish decreased during starvation, while the differences in mean weight before and after the starvation period were not significant in fish fed a level of 0.2% BW day^?1 and those fish fed to satiation. No mortality was recorded in either experiment. Results of this study indicate that a feeding rate of 1% BW day^?1 would be sufficient for commercial fish farming of YOY of this species to maintain them over winter. Also, to maintain fish weights and prevent weight loss in overwintering ponds, a feeding rate of around 0.3% BW day^?1 seems appropriate for hatcheries.     

Regulation of body weight and thermogenesis in seasonally acclimatized Brandt's voles (Microtus brandti)

Seasonal changes in an animal's morphology, physiology, and behavior are considered to be an adaptive strategy for survival and reproductive success. In the present study, we examined body weight and several behavioral, physiological, hormonal, and biochemical markers in seasonally acclimatized Brandt's voles (Microtus brandti) to test our hypothesis that Brandt's voles can decrease energy intake associated with decrease in body weight, body fat content, serum leptin level, and increasing thermogenesis in winter conditions. We found that the body weight of Brandt's voles was lowest in winter (December to February) and highest in spring and early summer (May to June). This seasonal variation in body weight was associated with changes in other markers examined. For example, the winter decrease in body weight was accompanied by increased energy intake and enhanced nonshivering thermogenesis (NST) as well as by decreased body fat mass and reduced levels of circulating leptin. Further, circulating levels of leptin were positively correlated with body weight and body fat mass, and negatively correlated with energy intake and uncoupling protein 1 contents. Together, these data do not support our hypothesis and suggest that leptin may be involved in this process and serve as a starvation signal in Brandt's voles.     

Assessing the role of body mass and sex on apparent adult survival in polygynous passerines: a case study of cetti's warblers in central Portugal

Adult survival, an important fitness component, is usually 1) lower in lighter individuals due to their reduced ability to survive winter conditions compared to heavier ones, especially in resident species at northern temperate latitudes and 2) lower in females compared with males due to higher reproductive costs incurred by females. In this paper, a capture–mark–recapture dataset of 649 cetti's warblers Cettia cetti ringed seasonally at two wetlands in central Portugal over an 11‐yr period (2000–2010) was modelled in a multi‐state framework to examine the influence of these individual covariates on apparent adult survival, while controlling for the presence of transient individuals in our study area. The probability of change in mass state (ψ_{Light→Heavy}, ψ_{Heavy→Light}) during the annual cycle was also estimated. Overall, birds survived better during spring–summer (breeding/moulting periods) compared with autumn–winter, but there was no effect of body mass on apparent adult survival probability. The modelling detected a significant interaction between sex and season, in which resident females survived better than resident males in spring–summer (?_RF= 0.857 ± 0.117 and ?_RM= 0.698 ± 0.181) while the opposite pattern was found in autumn–winter (?_RM= 0.440 ± 0.086 and ?_RF= 0.339 ± 0.084). In addition, cetti's warblers had a tendency to lose mass in spring–summer (ψ_{Heavy → Light}= 0.560 ± 0.063) and to regain mass in autumn–winter (ψ_{Light→Heavy}= 0.701 ± 0.069). This pattern of body mass change during the annual cycle may reflect energetic costs to reproduction and moulting, and/or a response to increased starvation risk during winter. High body mass, however, did not increase adult survival in this population presumably due to the relatively mild winter weather prevailing in central Portugal. Survival estimates are more likely to be explained by important ecological and behavioural differences between the two sexes in polygynous passerines. Our results highlight that studies aiming to identify the main factors shaping survival and individual fitness in polygynous species should be conducted during different phases of their annual cycle.     

Density-dependence vs. density-independence - linking reproductive allocation to population abundance and vegetation greenness

1. Recent studies have shown that optimal reproductive allocation depends on both climatic conditions and population density. We tested this hypothesis using six years of demographic data from eight reindeer (Rangifer tarandus) populations coupled with data on population abundance and vegetation greenness [measured using the Enhanced Vegetation Index (EVI)]. 2. Female spring body mass positively affected summer body mass gain, and lactating females were unable to compensate for harsh winters as efficiently as barren ones. Female spring body mass was highly sensitive to changes in population abundance and vegetation greenness and less dependent on previous autumn body mass and reproductive status. Lactating females were larger than barren females in the spring. Moreover, female autumn body mass was positively related to female autumn body mass and reproductive success and was not very sensitive to changes in vegetation greenness and population abundance. 3. Offspring autumn body mass was positively related to both maternal spring and autumn body mass, and as predicted from theory, offspring were more sensitive to changes in vegetation greenness and population abundance than adult females. A lagged cost of reproduction was present as larger females who were barren, the previous year produced larger offspring than equally sized females that successfully reproduced the previous year. 4. Reproductive success was negatively related to female autumn body mass and positively related to female spring body mass. Moreover, females who successfully reproduced the previous year experienced the highest reproductive success. The fact that negative density-dependence was only present for females that had successfully reproduced the previous year further support the hypothesis that reproduction is costly. 5. This study shows that female reindeer buffer their reproductive allocation according to expected winter conditions and that their buffering abilities were limited by population abundance and a lagged cost of reproduction and enhanced by vegetation greenness.     

Observational evidence of risk-sensitive reproductive allocation in a long-lived mammal

Organisms should adopt a risk-sensitive reproductive allocation when summer reproductive allocation competes with survival in the coming winter. This trade off is shown through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis on female reindeer in a population that has experienced a time period of dramatic increase in abundance. Environmental conditions during winter were fairly stable (with the exception of 1 year). We conclude that increased population abundance (perhaps in interaction with winter environmental conditions) could have represented a worsening of winter environmental conditions as both autumn offspring and spring female body mass decreased during the course of the study. Moreover, we found that the cost of reproduction was related to environmental conditions as: (1) autumn body mass was larger for barren than for lactating females, and this difference was temporally highly variable; (2) lactating females produced smaller offspring than barren ones in the following year; and (3) reproductive output (offspring size) decreased over time. We also found evidence of quality effects as lactating females had a higher reproductive success in the following year. In sum, a worsening of winter conditions lead to: (1) decreased reproductive output; (2) lowered autumn body mass for lactating females; and (3) increased body mass for barren females. Since females reduce their reproductive allocation as winter conditions becomes more severe, we conclude that reindeer have adopted a risk-sensitive reproductive allocation.     

Fattening strategies of wintering great tits support the optimal body mass hypothesis under conditions of extremely low ambient temperature

1.  In a seasonal environment, subcutaneous energy reserves of resident animals often increase in winter and decline again in summer reflecting gradual seasonal changes in their fattening strategies. We studied changes in body reserves of wintering great tits in relation to their dominance status under two contrasting temperature regimes to see whether individuals are capable of optimizing their body mass even under extreme environmental conditions.
2.  We predicted that dominant individuals will carry a lesser amount of body reserves than subordinate great tits under mild conditions and that the body reserves of the same dominant individuals will increase and exceed the amount of reserves of subordinates under conditions of extremely low ambient temperatures, when ambient temperature dropped down to −37 °C.
3.  The results confirmed the predictions showing that dominant great tits responded to the rising risk of starvation under low temperatures by increasing their body reserves and this was done at the expense of their safety.
4.  Removal experiments revealed that lower body reserves of subordinate flock members are due to the increased intraspecific competition for food under low ambient temperatures.
5.  Our results also showed that fattening strategies of great tits may change much quicker than previously considered, reflecting an adaptive role of winter fattening which is sensitive to changes in ambient temperatures.     

Fit females and fat polygynous males: seasonal body mass changes in the grey-headed flying fox

When females and males differ in their timing of maximum reproductive effort, this can result in sex-specific seasonal cycles in body mass. Such cycles are undoubtedly under strong selection, particularly in bats, where they affect flying ability. Flying foxes (Old World fruit bats, Pteropus spp.) are the largest mammals that can sustain powered flight and therefore face critical trade-offs in managing body reserves for reproduction, yet little is known about body mass dynamics in this group. I investigated body mass changes in relation to reproductive behaviour in a large colony of grey-headed flying foxes (Pteropus poliocephalus). In this polygynous mammal, females were predicted to maximise reproductive effort during lactation and males during the breeding season. As predicted, female body condition declined during the nursing period, but did not vary in relation to sexual activity. By contrast, males accumulated body reserves prior to the breeding season, but subsequently lost over 20% of their body mass on territory defence and courtship, and lost foraging opportunities as they also defended their day roost territories at night. Males in better condition had larger testes, particularly during territory establishment, prior to maximum sexual activity. Thus, the seasonality of female mass reflected the high metabolic load that lactation imposes on mothers. However, male mass followed a pattern akin to the "fatted male phenomenon", which is commonly observed in large polygynous mammals with seasonal reproduction, but not in bats. This shows the importance of body reserves for reproduction in flying foxes, despite their severe constraints on body mass.