Influence of ambient temperature on the phenology of the greater mouse-eared bat (Myotis myotis).

In order to assess the consequences of climate change and evaluate its impacts on wildlife, it is essential to do so on a species-specific level. It is assumed that changes in the ambient temperature influence energy consumption as well as food availability and thus foraging behavior, reproduction, survival, and therefore population dynamics in bats. Based on this assumption, the present study aims to gain insights into the roosting and breeding behavior of the greater mouse-eared bat (Myotis myotis) in relation to changes of the ambient temperature. For this purpose, we investigated the effect of ambient temperature on the phenology of the greater mouse-eared bat by using activity data of the bats collected using light barriers at the maternity roosts. The light barrier used in this study is a system that detects the interruption of two light beams, for example, by a flying bat, and displays it as an electrical signal. The investigations have shown that the higher the winter temperatures, the earlier the greater mouse-eared bats returned to the roosts to form the maternity colony; however, this was only true for ambient temperatures below 0.5°C,birth season started earlier at higher spring temperatures,the dissolution of maternity roosts occurred earlier with earlier birth season and at higher ambient temperatures during lactation. The results revealed that ambient temperature has an influence on the phenology of the greater mouse-eared bat. Depending on the respective life history stage, an increase in ambient temperature can have a positive or negative effect on the fitness of the animals. In recent years, mild winters have been recorded more frequently, which can have an influence on the behavior of bats. Warm winters within certain limits seem to lead to an earlier formation of the maternity colony, which can be positive or negative for the bats depending on persistent weather conditions and thus insect availability. In the course of climate change, we can also expect earlier spring events and an increase in spring temperature, as well as hot spells in summer. These warm springs and summers seem to lead to an earlier beginning of births, a faster development of the juveniles and an earlier dissolution of the maternity roost. An advance of reproductive activities can be assumed to increase the chance to survive the following winter in both mothers and their young, as they have more time to build up sufficient energy reserves for hibernation before winter starts. Due to the climatic changes, phenological changes of the bats be expected. This study highlights that in order to understand the impact of climate change on biodiversity, it is necessary to investigate in detail effects on a species-specific level and also to consider direct and indirect effects of ambient temperature on different life history stages.

Season and reproductive activity influence cortisol levels in the Malagasy primate Lepilemur edwardsi.

Throughout the year, wild animals are exposed to a variety of challenges such as changing environmental conditions and reproductive activity. These challenges may affect their stress hormone levels for varying durations and in varying intensities and impacts. Measurements of the glucocorticoid hormone cortisol in the hair of mammals are considered a good biomarker for measuring physiological stress and are increasingly used to evaluate stress hormone levels of wild animals. Here, we examined the influence of season, reproductive activity, sex, as well as body condition on hair cortisol concentrations (HCC) in Lepilemur edwardsi, a small Malagasy primate species. L. edwardsi lives in the seasonal dry forests of western Madagascar, which are characterized by a strongly changing resource availability throughout the year. We hypothesized that these seasonal changes of resource availability and additionally the reproductive cycle of this species would influence HCC of L. edwardsi. Results revealed that hair cortisol concentration of females did not change seasonally or with the reproductive cycle. However, we found a significant increase of hair cortisol levels in males from the early wet season during the early dry season (mating season). This increase is presumably due to changed behavior during the mating season, as sportive lemurs travel more and show aggressive behavior during this time of the year. This behavior is energy-costly and stressful, and presumably leads to elevated HCC. As elevated cortisol levels may impair immune function, L. edwardsi males might also be more susceptible to parasites and diseases, which is unfavorable in particular during a period of low resource availability (dry season).

Bridging environment, physiology and life history: Stress hormones in a small hibernator.

Our knowledge of the perception of stress and its implications for animals in the wild is limited, especially in regard to mammals. The aim of this study was therefore to identify sex specific effects of reproductive activity, body mass, food availability and hibernation on stress hormone levels in the edible dormouse (Glis glis), a small mammalian hibernator. Results of our study reveal that reproductive activity and pre-hibernation fattening were associated with high cortisol levels in both sexes. During the mating season, in particular individuals with low body masses had higher stress levels. Elevated levels of cortisol during pre-hibernation fattening were even higher in females that had formerly invested into reproduction compared to non-reproductive females. Previously observed impairments on health parameters and reduced survival rates associated with reproduction emphasize the functional relevance of high stress hormone levels for fitness. Prolonged food limitation, however, did not affect stress levels demonstrating the ability of dormice to predict and cope with food restriction.

Flexibility is the key: metabolic and thermoregulatory behaviour in a small endotherm.

Hibernation is the most effective way to reduce thermoregulatory costs during periods of unfavourable environmental conditions. In preparation to hibernation, fat-storing hibernators accumulate large quantities of body fat, which increases their locomotor costs and also the risk of predation. As a consequence, there should be a strong selective pressure to restrict pre-hibernation fattening to a short-time period before the onset of hibernation. The edible dormouse (Glis glis) is characterized by having adapted its whole life history to the irregularly occurring mast-seeding pattern of the European beech (Fagus sylvaticus). Thus, the question arises how this small endotherm copes with huge differences in food availability between years. Therefore, we investigated body mass and thermal energetics of edible dormice during high and low food years. Our results demonstrate that during periods of low food availability, edible dormice enter an energy-saving mode with reduced body temperature (Tb) and resting metabolic rate (RMR), and high torpor frequencies. During irregularly occurring short events of high food availability in mast years, however, Tb was higher, torpor did not occur, and RMR was drastically elevated possibly due to an enlarged digestive tract and the heat increment of feeding associated with a dietary switch to high-quality food and an increase in the amount of food ingested. This physiological flexibility allows edible dormice to efficiently accumulate body fat reserves under extremely different situations of food availability.

Life history written in blood: erythrocyte parameters in a small hibernator, the edible dormouse.

The oxygen delivery system is one major determinant of the performance of vertebrates and responds sensitively to a variety of internal and environmental factors. To understand physiological mechanisms underlying variations of fitness, we investigated effects of demanding conditions associated with certain life-history events, food availability, and population density on the oxygen delivery system in free-ranging edible dormice (Glis glis). We sampled blood (n = 248) and urine (n = 319), performed an erythrocyte haemogram and visually determined the presence of haemoglobinuria. Reproduction leads to increased mortality in edible dormice and our study now reveals severe haematological impairments during reproduction that were associated with nutrient and energy deficits and stress. These effects were even more pronounced in subsequent reproductive years, indicating prolonged physiological impairment. Under limited food availability, the rate of erythrocyte generation was reduced. This seems to be part of an energy saving strategy instead of representing a poor body condition as survival probability in this species is high in years of low food availability. A high prevalence ratio of haemoglobinuria (up to 85%) at the end of the active season indicated amplified erythrocyte destruction through haemolysis. This may be the result of a preparative mechanism to avoid massive oxidative damage during the long hibernation period. Most ecophysiological studies so far focus on single erythrocyte parameters on a short time scale, which could be misleading. Our results clearly highlight that a wide-array RBC approach is a powerful tool for investigating mechanisms underlying physiological performance and fitness, also for other vertebrate taxa.

It takes two to tango: Phagocyte and lymphocyte numbers in a small mammalian hibernator.

Immunity is energetically costly and competes for resources with other physiological body functions, which may result in trade-offs that impair fitness during demanding situations. Endocrine mediators, particularly stress hormones, play a central role in these relationships and directly impact leukocyte differentials. To determine the effects of external stressors, energetic restraints and competing physiological functions on immune parameters and their relevance for fitness, we investigated leukocyte profiles during the active season of a small obligate hibernator, the edible dormouse (Glis glis), in five different study sites in south-western Germany. The highly synchronized yearly cycle of this species and the close adaptation of its life history to the irregular abundance of food resources provide a natural experiment to elucidate mechanisms underlying variations in fitness parameters. In contrast to previous studies on hibernators, that showed an immediate recovery of all leukocyte subtypes upon emergence, our study revealed that hibernation results in depleted phagocyte (neutrophils and monocytes) stores that recovered only slowly. As the phenomenon of low phagocyte counts was even more pronounced at the beginning of a low food year and primarily immature neutrophils were present in the blood upon emergence, preparatory mechanisms seem to determine the regeneration of phagocytes before hibernation is terminated. Surprisingly, the recovery of phagocytes thereafter took several weeks, presumably due to energetic restrictions. This impaired first line of defense coincides with lowest survival probabilities during the annual cycle of our study species. Reduced survival could furthermore be linked to drastic increases in the P/L ratio (phagocytes/lymphocytes), an indicator of physiological stress, during reproduction. On the other hand, moderate augmentations in the P/L ratio occurred during periods of low food availability and were associated with increased survival, but reproductive failure. In this case, the stress response probably represents an adaptive reaction that contributes to survival by activating energy resources. In contrast to our expectation, we could not detect an amplification of stress through high population densities. Summarized, results of our study clearly reveal that the leukocyte picture of active edible dormice responds sensitively to physiological conditions associated with hibernation, reproductive activity and food availability and can be linked to fitness parameters such as survival. Thus edible dormice represent an excellent model organism to investigate regulatory mechanisms of the immune system under natural conditions.

The vector tick Ixodes ricinus feeding on an arboreal rodent-the edible dormouse Glis glis.

The reservoir competence and long life expectancy of edible dormice, Glis glis, suggest that they serve as efficient reservoir hosts for Lyme disease (LD) spirochetes. Their arboreality, however, may reduce the probability to encounter sufficient questing Ixodes ricinus ticks to acquire and perpetuate LD spirochetes. To define the potential role of this small arboreal hibernator in the transmission cycle of LD spirochetes, we examined their rate and density of infestation with subadult ticks throughout the season of activity. Of the 1081 edible dormice that we captured at five study sites in Southern Germany and inspected for ticks at 2946 capture occasions, 26 % were infested with at least one and as many as 26 subadult ticks on their ear pinnae. The distribution of ticks feeding on edible dormice was highly aggregated. Although only few individuals harbored nymphal ticks soon after their emergence from hibernation, the rate of nymphal infestation increased steadily throughout the season and reached about 35 % in September. Dormice inhabiting a site with few conspecifics seemed more likely to be infested by numerous ticks, particularly nymphs, than those individuals living in densely populated sites. Male dormice were more likely to be parasitized by numerous nymphs than were females, independent of their age and body mass. Our observation that season, population density, and sex affect the rates of ticks feeding on edible dormice suggests that the contribution of edible dormice to the transmission cycle of LD spirochetes depends mainly on their ranging behavior and level of activity.

Ways to measure body temperature in the field.

Body temperature (Tb) represents one of the key parameters in ecophysiological studies with focus on energy saving strategies. In this study we therefore comparatively evaluated the usefulness of two types of temperature-sensitive passive transponders (LifeChips and IPTT-300) and one data logger (iButton, DS1922L) mounted onto a collar to measure Tb in the field. First we tested the accuracy of all three devices in a water bath with water temperature ranging from 0 to 40°C. Second, we evaluated the usefulness of the LifeChips and the modified iButtons for measuring Tb of small heterothermic mammals under field conditions. For this work we subcutaneously implanted 14 male edible dormice (Glis glis) with transponders, and equipped another 14 males with data loggers to simultaneously record Tb and oxygen consumption with a portable oxygen analyzer (Oxbox). In one individual we recorded Tb with both devices and analyzed recorded Tb patterns. LifeChips are able to measure temperature within the smallest range from 25 to 40°C with an accuracy of 0.07±0.12°C. IPTT-300 transponders measured temperature between 10 and 40°C, but accuracy decreased considerably at values below 30°C, with maximal deviations of nearly 7°C. An individual calibration of each transponder is therefore needed, before using it at low Tbs. The accuracy of the data logger was comparatively good (0.12±0.25°C) and stable over the whole temperature range tested (0-40°C). In all three devices, the repeatability of measurements was high. LifeChip transponders as well as modified iButtons measured Tb reliably under field conditions. Simultaneous Tb-recordings in one edible dormouse with an implanted LifeChip and a collar-mounted iButton revealed that values of both measurements were closely correlated. Taken together, we conclude that implanted temperature-sensitive transponders represent an appropriate and largely non-invasive method to measure Tb also under field conditions.

Seasonal prevalence of Lyme disease spirochetes in a heterothermic mammal, the edible dormouse (Glis glis).

In Europe, dormice serve as competent reservoir hosts for particular genospecies of the tick-borne agent of Lyme disease (LD) and seem to support them more efficiently than do mice or voles. The longevity of edible dormice (Glis glis) and their attractiveness for ticks may result in a predominance of LD spirochetes in ticks questing in dormouse habitats. To investigate the role of edible dormice in the transmission cycle of LD spirochetes, we sampled skin tissue from the ear pinnae of dormice inhabiting five different study sites in south western Germany. Of 501 edible dormice, 12.6% harbored DNA of LD spirochetes. Edible dormice were infected most frequently with the pathogenic LD spirochete Borrelia afzelii. The DNA of B. garinii and B. bavariensis was detected in ca. 0.5% of the examined individuals. No spirochetal DNA was detectable in the skin of edible dormice until July, 6 weeks after they generally start to emerge from their obligate hibernation. Thereafter, the prevalence of spirochetal DNA in edible dormice increased during the remaining period of their 4 to 5 months of activity, reaching nearly 40% in September. Males were more than four times more likely to harbor LD spirochetes than females, and yearlings were almost twice more likely to be infected than adults. The seasonality of the prevalence of LD spirochetes in edible dormice was pronounced and may affect their role as a reservoir host in respect to other hosts.

Genetic consequences of forest fragmentation for a highly specialized arboreal mammal--the edible dormouse.

Habitat loss and fragmentation represent the most serious extinction threats for many species and have been demonstrated to be especially detrimental for mammals. Particularly, highly specialized species with low dispersal abilities will encounter a high risk of extinction in fragmented landscapes. Here we studied the edible dormouse (Glis glis), a small arboreal mammal that is distributed throughout Central Europe, where forests are mostly fragmented at different spatial scales. The aim of this study was to investigate the effect of habitat fragmentation on genetic population structures using the example of edible dormouse populations inhabiting forest fragments in south western Germany. We genotyped 380 adult individuals captured between 2001 and 2009 in four different forest fragments and one large continuous forest using 14 species-specific microsatellites. We hypothesised, that populations in small forest patches have a lower genetic diversity and are more isolated compared to populations living in continuous forests. In accordance with our expectations we found that dormice inhabiting forest fragments were isolated from each other. Furthermore, their genetic population structure was more unstable over the study period than in the large continuous forest. Even though we could not detect lower genetic variability within individuals inhabiting forest fragments, strong genetic isolation and an overall high risk to mate with close relatives might be precursors to a reduced genetic variability and the onset of inbreeding depression. Results of this study highlight that connectivity among habitat fragments can already be strongly hampered before genetic erosion within small and isolated populations becomes evident.

Survival rates in a small hibernator, the edible dormouse: a comparison across Europe.

Understanding how local environmental factors lead to temporal variability of vital rates and to plasticity of life history tactics is one of the central questions in population ecology. We used long-term capture-recapture data from five populations of a small hibernating rodent, the edible dormouse Glis glis, collected over a large geographical range across Europe, to determine and analyze both seasonal patterns of local survival and their relation to reproductive activity. In all populations studied, survival was lowest in early summer, higher in late summer and highest during hibernation in winter. In reproductive years survival was always lower than in non-reproductive years, and females had higher survival rates than males. Very high survival rates during winter indicate that edible dormice rarely die from starvation due to insufficient energy reserves during the hibernation period. Increased mortality in early summer was most likely caused by high predation risk and unmet energy demands. Those effects have probably an even stronger impact in reproductive years, in which dormice were more active. Although these patterns could be found in all areas, there were also considerable differences in average survival rates, with resulting differences in mean lifetime reproductive success between populations. Our results suggest that edible dormice have adapted their life history strategies to maximize lifetime reproductive success depending on the area specific frequency of seeding events of trees producing energy-rich seeds.

Behavioural and physiological consequences of male reproductive trade-offs in edible dormice (Glis glis).

Testosterone mediates male reproductive trade-offs in vertebrates including mammals. In male edible dormice (Glis glis), reproductivity linked to high levels of testosterone reduces their ability to express torpor, which may be expected to dramatically increase thermoregulatory costs. Aims of this study were therefore to analyse behavioural and physiological consequences of reproductive activity in male edible dormice under ecologically and evolutionary relevant conditions in the field. As we frequently encountered sleeping groups in the field, we hypothesized that social thermoregulation should be an important measure to reduce energy expenditure especially in sexually active male edible dormice. Our results revealed that the occurrence of sleeping groups was negatively influenced by male body mass but not by reproductive status or ambient temperature. In reproductive as in non-reproductive males, the number of individuals huddling together was negatively influenced by their body mass. Thus in general males with a high body mass were sitting in smaller groups than males with a low body mass. However, in reproductive males group size was further negatively affected by ambient temperature and positively by testes size. Thus breeders formed larger sleeping groups at lower ambient temperatures and males with larger testes were found in larger groups than males with smaller testes. Measurements of oxygen consumption demonstrated that grouping behaviour represents an efficient strategy to reduce energy expenditure in edible dormice as it reduced energy requirements by almost 40%. In summary, results of this field study showcase how sexually active male edible dormice may, through behavioural adjustment, counterbalance high thermoregulatory costs associated with reproductive activity.

Functional associations of similar MHC alleles and shared parasite species in two sympatric lemurs.

Several recent studies of animals in their natural surroundings found evidence for effects of certain major histocompatibility complex (MHC) immune gene alleles on the parasite load. However, in multi-infected individuals the particular selection pressure exerted by specific parasites has rarely been explored. In this study we took advantage of the parasitological and genetic data of two previously investigated Malagasy lemur species (Cheirogaleus medius and Microcebus murinus). We investigated whether the two sympatric and ecologically similar primates are infected by similar parasite species and explored if certain parasites are associated with particular MHC alleles. Our study revealed that most of the parasite egg morphotypes were found in both hosts. In each lemur species we identified one MHC allele which was positively associated with Ascaris-infection. Interestingly, these MHC alleles were very similar to each other but differed from all other investigated MHC alleles in an amino acid substitution in a putative functional important antigen binding site. Thus, our study gives first intriguing evidence for a direct connection between certain antigen binding sites of MHC molecules with a particular parasite in two wild primate populations. This may indicate that indeed certain parasites exert direct selective pressure on the MHC of wild living hosts.

Is energy supply the trigger for reproductive activity in male edible dormice (Glis glis)?

In edible dormice (Glis glis) reproduction is synchronised with the intermittent masting of the European beech (Fagus sylvatica). In years of mast failure dormouse males seem to anticipate future low food availability and fail to develop functional testes. We hypothesised that the availability of high-quality food is linked to male reproductive capacity, because of high male energetic demands during gonad maturation. We therefore evaluated the relationship between beech seed production and male reproductivity in the field between 1993 and 2005. In order to know whether the energy content of the food as such triggers sexual capacity, we supplemented high-quality food in the field for 3 years and investigated reproductive output, reproductive capacity, and body mass changes. Results revealed that male reproductive capacity was positively linked with beech seed production. Body mass changes of reference males during the high reproductive year further revealed high energetic demands of male reproduction, which were counter balanced in food-supplemented males. However, in contrast to our assumptions, artificial food supply during a year of mast failure failed to evoke high reproductivity in edible dormice. The availability of high-quality food can therefore be ruled out from being the primary trigger for sexual activity in male edible dormice.

Influences of the feeding ecology on body mass and possible implications for reproduction in the edible dormouse (Glis glis).

The edible dormouse (Glis glis) is a small rodent and an obligate hibernator. Dormice undergo strong fluctuations of reproductive output during years that seem to be timed to coincide with future food supply. This behaviour enables them to avoid producing young that will starve with a high probability due to food shortage, and to increase their lifetime reproductive success. Aims of this study were to elucidate the extent to which feeding ecology in the edible dormouse has an impact on body mass and the fatty acid (FA) pattern of the white adipose tissue (WAT) before and after hibernation, which in turn might influence reproductive status in spring. Dormice show strong seasonal fluctuations of the body mass, which is reduced by one third during hibernation. Body mass and its changes depend on autumnal food availability as well as on the dietary FA pattern. During the pre-hibernation fattening period, dormice eat lipid rich food with a high content of linoleic acid. During hibernation, linoleic acid content is slightly but significantly reduced and body mass loss during winter is negatively correlated with the pre-hibernation linoleic acid content in the WAT. No relation between reproductive status and body mass, body condition or the FAs pattern of the WAT could be detected. However, in a year of high reproduction, dormice commence the shift to seed eating earlier than in a year of low reproduction. These seeds could be either a predictor for future food supply in autumn, or represent a high-energy food compensating high energetic costs of sexual activity in male edible dormice.

Energetic constraints on sexual activity in the male edible dormouse (Glis glis).

The aim of this study was to examine to what extent reproductive activity in male edible dormice (Glis glis) might be energetically constrained. Demographic data, morphometric data, and oral body temperature (T(or)) measurements were collected in two study areas between 1993 and 2002 in southwest Germany and combined with subcutaneous body temperature (T(sc)) registrations of captive dormice. T(sc) measurements were collected directly after emergence from hibernation (June) until the end of the mating season (July). Wild edible dormice showed strong fluctuations in their reproductive output between years. Not all males were sexually active each year and the number of litters born was positively correlated with the number of sexually active males, which suggests that sexual activity in males is constrained and in turn limits reproductive success. A comparison of the T(or) of sexually quiescent and active males revealed that sexually quiescent males had significantly lower T(or) (median: 28.8 degrees C; 25/75% quartiles: 16.4/31.0; n=31) than sexually active males (median: 34.2 degrees C; 25/75% quartiles: 32.0/35.6; n=156). Body condition of sexually active and quiescent males was not different after emergence from hibernation. However, sexually active males showed a significant reduction in their body condition between June and July, the time of mating, while body condition of sexually quiescent males remained constant. Continuous T(sc) registrations in captive sexually active male dormice showed strong circadian T(sc) fluctuations. Even though daily torpor bouts with T(sc) below 20 degrees C occurred in these males, most of the time T(sc) fluctuated above 30 degrees C, which is known as the critical body temperature threshold above which testes maturation can take place in this species. These results demonstrate that male dormice incur high costs due to sexual activity and that thermoregulation is determined by a trade-off between energetic savings and reproductive activity.

Costs and potential benefits of parental care in the nocturnal fat-tailed dwarf lemur (Cheirogaleus medius).

Parental care should evolve only if fitness benefits exceed fitness costs. In Cheirogaleus medius, a small nocturnal lemur of western Madagascar, we found the peculiar situation of considerable parental care by both sexes, combined with an extremely high rate of extra-pair young (EPY). In this paper, we try to elucidate the costs and benefits of parental care in C. medius, and we discuss hypotheses as to why males might actively participate in raising young, especially with regard to the high rate of EPY. The study was carried out in the dry deciduous forest of western Madagascar between 1996 and 2001. The most important components of parental care in C. medius are babysitting and guarding of the young. Observational data suggest that thermoregulatory demands during the first days after birth and defence from predators may be the driving factors selecting for this infant care behaviour. The reproducing individuals of both sexes incur considerable energetic costs by this behaviour, resulting in reduced fat stores and body condition in comparison to non-reproducing individuals. The reason why males also care for EPY remains elusive. We propose that males might not be able to detect individual relatedness and that they would jeopardise the survival of their own young if they gave no parental care. Alternatively, they might gain advantages other than direct fitness from raising EPY, for example if caring behaviour increases their chances in further reproduction, or if EPY are fathered by close kin.

Sex-specific recombination rates in Parus major and P. caeruleus, an exception to Huxley's rule.

Huxley's rule predicts lower recombination rates in the heterogametic sex than in the homogametic one. The genotyping of Parus major and P. caeruleus families at 8 microsatellite and 4 enzyme loci yielded contradicting data. Significant genotypic disequilibrium was observed between esterase-1, esterase-2 and esterase-3 in adults of P. major and between esterase-2/esterase-3 and esterase-2/microsatellite PK-12 in P. caeruleus. Support comes from linkage analyses of nuclear families. In P. major, the recombination rate of esterase-2/esterase -3 in males is significantly lower than in females (theta(male) = 0.076, theta(female) = 0.145). The opposite is found for the recombination rates of esterase-1/esterase-2 and esterase-2/esterase-3 in P. caeruleus (EST-1/EST-2: theta(female) = 0.218, theta(male) = 0.5, EST-2/EST-3: theta(female) = 0.109, theta(male) = 0.194). We conclude that the basis of differences in recombination rates cannot be heterogamety, per se, but must have multiple genetic causes including chromosomal rearrangments that have evolved after the cladogenesis of the two species.

Feeding ecology of the hibernating primate Cheirogaleus medius: how does it get so fat?

The fat-tailed dwarf lemur (Cheirogaleus medius; Cheirogaleidae) lives in the tropical forests of Madagascar. It is exceptional among primates, because this small nocturnal lemur hibernates during the dry season from the end of April until October. This field study was carried out to assess seasonal changes in body mass and behavioral traits (locomotor activity, feeding ecology) of C. medius that enable this species to accumulate large quantities of fat within weeks. Feeding observations of 36 radio-collared individuals were combined with a mark-recapture study in western Madagascar during the rainy seasons from 1995 to 1998. Pulp of fruit species not consumed, as well as fruit species eaten by C. medius was analyzed for lipid and sugar contents. Body mass changes of recaptured individuals revealed that C. medius may double its body mass within a few weeks. Most of the increase in body mass took place between March and April after infants had been weaned and directly before the onset of hibernation. Locomotor activity was significantly reduced during the period of body mass gain. During the entire study, fruits, nectar, and flowers of 34 different plant species comprised about 80% of the diet, supplemented by a seasonally varying intake of animal prey. Lipid contents of consumed fruits were comparatively low throughout the study period. No difference in lipid content was detectable between fruit species consumed and not consumed by C. medius. During prehibernation fattening, C. medius consumed mostly fruit species with a high sugar content, significantly higher than that of fruit species that were not consumed. The results suggest that a reduction in locomotor activity as well as a preference for fruit species with a high sugar content enable C. medius to deposit large quantities of fat in a few weeks.