Seasonal dynamics of agonistic behavior and hormones in an ex situ all‐male colony of large flying foxes

Large flying foxes (Pteropus vampyrus) are a socially complex species. In situ colonies typically comprise thousands of individuals in small harems of one male to many females. In ex situ environments, all‐male colonies are becoming more common due to a surplus of males in the population. There is limited information describing the hormonal and behavioral patterns of all‐male colonies during the breeding season. We assessed seasonal changes in hormones and behavior in an all‐male colony of 12 large flying foxes at Disney's Animal Kingdom^®. We validated hormone assays using morning urine and fecal samples to assess seasonal changes in excreted immunoreactive testosterone and glucocorticoid metabolites. We collected behavior data using an all‐occurrence method, recording agonistic behaviors related to territorial defense (hooking, biting, wing flexing, vocalizing, and wrestling), and sexual behavior (mounting and frontal grabbing). Results indicated that (i) we could reliably measure testosterone and glucocorticoid metabolites concentrations from fecal and urine samples collected from individual bats; (ii) there were distinct relationships between changes in levels of agonism and hormone concentrations throughout the year; and (iii) three agonistic behaviors (chasing, wrestling, and open‐mouth threat) peaked prior to the increase in testosterone and glucocorticoid hormones measured during the breeding season. These three behaviors could potentially be used as early indicators to signal the onset of the breeding season and allow time to implement ex situ management changes to reduce the incidence of agonism between individuals.     

Impact of body reserves on energy expenditure, water flux, and mating success in breeding male northern elephant seals

In capital breeders, individual differences in body size and condition can impact mating effort and success. In addition to the collateral advantages of large body size in competition, large nutrient reserves may offer advantages in endurance rivalry and enable the high rates of energy expenditure associated with mating success. We examined the impacts of body reserves and dominance rank on energy expenditure, water flux, mating success, and breeding tenure in the adult male northern elephant seal, a polygynous, capital breeder. Adult males expended energy at a rate of 159 ± 49 MJ d (-1), which is equivalent to 3.1 times the standard metabolic rate predicted by Kleiber's equation. Despite high rates of energy expenditure and a long fasting duration, males spared lean tissue effectively, deriving a mean of 7% of their metabolism from protein catabolism. Body composition had a strong impact on the ability to spare lean tissue during breeding. When controlling for body size, energy expenditure, depletion of blubber reserves, and water efflux were significantly greater in alpha males than in subordinate males. Large body size was associated with increased reproductive effort, tenure on shore, dominance rank, and reproductive success. Terrestrial locomotion and topography appeared to strongly influence energy expenditure. Comparisons with conspecific females suggest greater total seasonal reproductive effort in male northern elephant seals when controlling for the effects of body mass. In polygynous capital breeding systems, male effort may be strongly influenced by physiological state and exceed that of females.     

Homosexual Fellatio: Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon

A recent focus of interest has been on the functional significance of genital licking (fellatio and cunnilingus) in relation to sexual selection in Pteropodid bats. In the present paper, a form of fellatio in wild Bonin flying foxes, Pteropus pselaphon, performed between adult males has been reported. During the mating season, adult flying foxes roost in same-sex groups, forming ball-shaped clusters which provide warmth. The female clusters may also contain a few males. Unassociated with allogrooming, same-sex genital licking occurred among males in the all male clusters. As such, male-male fellatio can be considered as homosexual behavior, two functional explanations could account for this behavior; the social bonding and the social tension regulation hypotheses suggested in a previous review. Given that neither the simpler alternative that in all male groups such fellatio may represent misdirected sexual behavior, nor the two previously proposed functional hypotheses were supported by the data, I propose another functional hypothesis. Homosexual fellatio in this species could help males solve inconsistent situations in the roost when there are conflicts between cooperative behavior for social thermoregulation and competition for mating.     

Costs of reproduction in male lizards, Sceloporus virgatus

Models of life history evolution typically assume a balance between the benefits of current reproductive activity and the costs to future reproductive success or survivorship, but empirical studies often find positive correlations between such components of fitness in undisturbed animal populations. I examined possible survivorship costs of reproduction in free-ranging male lizards, Sceloporus virgatus , and found that males with low levels of mating success were less likely to survive to the following breeding season. I also investigated two possible indicators of reproductive effort, increase in ectoparasite load and decrease in body weight during the breeding season. Levels of parasitism with trombiculid mites at the end of the breeding season were not associated with any measure of fitness or body condition (mating success, survivorship to the following year, relative weight loss). Yearling males (which have low levels of mating success) usually gained weight during the breeding season, while older males generally lost weight during this period. This suggests that young males may have postponed reproduction in favor of body growth and that seasonal weight loss of older males might reflect reproductive effort. Within the group of older males, individuals with the highest levels of mating success did not have high levels of either weight loss or mortality. Mate guarding behavior, an alternative to the aggressive territorial behavior typical of many lizard species, may allow certain males to obtain mates without expending large amounts of energy or exposing themselves to great mortality risks.     

The hormonal and behavioral response to group formation, seasonal changes, and restraint stress in the highly social Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus) (Chiroptera: Pteropodidae)

This study examined behavioral and physiological responses (changes in inter-animal spacing, total glucocorticoids, testosterone, and body mass) to the formation of breeding and same-sex groups in two bat species, the socially gregarious Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus). We hypothesized that social instability, especially in the breeding groups and especially in P. vampyrus, would result in elevated glucocorticoids and that social facilitation of breeding and/or male-male competition would result in persistently higher levels of testosterone in breeding males. Seasonal rhythms in all measures were also predicted, and the glucocorticoid stress response was expected to vary by sex, season, and group type. Nearly all animals responded to group formation with elevated glucocorticoids, but, for breeding animals (especially aggressive male P. vampyrus), these responses persisted over time. In both species, breeding group formation resulted in elevated testosterone in males. Glucocorticoids, testosterone, testes volume, and body mass generally peaked in the breeding season in males (late summer and early autumn), but the seasonal glucocorticoid peak in females occurred in late winter and early spring. All animals responded to restraint stress with elevations in glucocorticoids that largely did not differ by sex, time of year, reproductive condition, group type, or, in lactating females, the presence of her pup. Changes in both behavior and physiology were more evident in P. vampyrus than in P. pumilus, and we believe that their underlying social differences influenced their responses to group formation and to the changing seasonal environment.     

Pollination by the locally endangered island flying fox (Pteropus hypomelanus) enhances fruit production of the economically important durian (Durio zibethinus)

Fruit bats provide valuable pollination services to humans through a unique coevolutionary relationship with chiropterophilous plants. However, chiropterophily in the Old World and the pollination roles of large bats, such as flying foxes (Pteropus spp., Acerodon spp., Desmalopex spp.), are still poorly understood and require further elucidation. Efforts to protect these bats have been hampered by a lack of basic quantitative information on their role as ecosystem service providers. Here, we investigate the role of the locally endangered island flying fox Pteropus hypomelanus in the pollination ecology of durian (Durio zibethinus), an economically important crop in Southeast Asia. On Tioman Island, Peninsular Malaysia, we deployed 19 stations of paired infrared camera and video traps across varying heights at four individual flowering trees in a durian orchard. We detected at least nine species of animal visitors, but only bats had mutualistic interactions with durian flowers. There was a clear vertical stratification in the feeding niches of flying foxes and nectar bats, with flying foxes feeding at greater heights in the trees. Flying foxes had a positive effect on mature fruit set and therefore serve as important pollinators for durian trees. As such, semi‐wild durian trees—particularly tall ones—may be dependent on flying foxes for enhancing reproductive success. Our study is the first to quantify the role of flying foxes in durian pollination, demonstrating that these giant fruit bats may have far more important ecological, evolutionary, and economic roles than previously thought. This has important implications and can aid efforts to promote flying fox conservation, especially in Southeast Asian countries.     

Regulation by Photoperiod of Seasonal Changes in Body Mass and Reproductive Function in Gray Mouse Lemurs (Microcebus murinus): Differential Responses by Sex

Microcebus murinus exhibits highly seasonal biological rhythms to cope with extreme seasonality in availability of resources. To study the role of daylength on seasonal changes in body mass and reproductive function, we exposed male and female gray mouse lemurs to natural, constant, or alternating light cycles for 2 years under constant environmental conditions. When exposed to either constant short (SD: 10 h light/day), long (LD: 14 h light/day), or intermediate (ID: 12 h light/day) daylength, males and females maintained a constant body mass with no spontaneous cyclic variation. We only observed typical seasonal body mass changes in subjects exposed to alternating periods of SD and LD, the weight gain being triggered by SD, whereas weight loss occurred under LD. Reproductive activity in females proceeded from an endogenous rhythm that was expressed under constant daylengths. In contrast, changes in reproductive activity in males depended on daylength variation. In both sexes, SD and LD have direct inhibitory or stimulatory effects on reproductive activity. In females, daylength regulates breeding season by synchronizing an endogenous sexual rhythm with the season, whereas in males, the perception of a critical photoperiod is used to determine the subsequent onset or arrest of their breeding season. These sexual differences in the effect of daylength could be related to sex-specific differences in reproductive constraints.     

Changes in body mass and feeding behaviour in male harbour seals, Phoca vitulina, in relation to female reproductive status

Loss of body mass has been used as an index of the cost of reproductive effort in mammals. We studied changes in body mass of male harbour seals, Phoca vitulina , during the breeding season on Sable Island, Nova Scotia, Canada. Individually marked subadult ( n = 21) and adult ( n = 22) males were captured at approximately six-day intervals throughout the breeding season. Adult males weighed an average of 108 kg ± 5·6 (S.E.) at initial capture, whereas subadults weighed an average of 76 ± 6·6 kg. The rate of mass loss by adult males did not differ from zero during the pre-mating period (i.e. the period without receptive females), but differed significantly from zero (-0·91 ± 0·007 kg/day) during the mating period (i.e. the period with receptive females). By contrast, the rate of mass change of subadults did not differ from zero in either period, indicating that food availability was unlikely to be responsible for the observed changes in adults. Adult males lost up to 24% of body mass during the breeding season. Examination of sera for the presence of chylomicrons (i.e. evidence of recent feeding) also suggested that adults stopped feeding during the mating period, while subadults did not. These results suggest that reproduction represents a significant energetic cost to adult male harbour seals.     

Reproductive strategies and the influence of date of birth on growth and sexual development of an aseasonally-breeding ungulate: Reeves' muntjac (Muntiacus reevesi)

Unlike most species of deer, Reeves' muntjac does not have a seasonal reproductive cycle. Births are equally distributed throughout the year and, irrespective of season of birth, females start breeding when they have reached a minimum body weight of about 10 kg. A significantly smaller proportion of females is born in the autumn/winter than the spring/summer, but there is no effect of maternal age or condition on foetal sex ratios for first or subsequent pregnancies. Whilst aseasonal breeding increases female productivity, males are only able to hold territories encompassing the ranges of a number of does for a relatively short period of time. It is argued that for muntjac there is much less inter-sexual variation in lifetime reproductive success than for seasonally-breeding polygynous cervids. Hence, although muntjac are sexually dimorphic and polygynous, females do not invest preferentially in male offspring.
Young males can be fertile from 36 weeks of age, when their first antlers are still in velvet. Whilst season of birth has no effect on the rate of sexual development in females, it does for males, with autumn-born male fawns attaining sexual maturity earliest. This accelerated development occurs in the period following independence from the mother, and there is no evidence of a maternal cost in producing male fawns. Since the canine tusks are the main weapons for intra-sexual conflict, there could be clear gains in terms of reproductive success for males that attain sexual maturity, adult weight and adult tusk size quickly, even though their first antlers are much smaller than those of older bucks. The hypothesis is presented that, for muntjac, the majority of a male's lifetime reproductive success is achieved relatively early in life when the canine tusks are in pristine condition, and that bucks lose their territories, and associated access to oestrous does, once the canines are broken.     

Seasonal changes in testicular size, plasma testosterone concentration and body weight in captive flying foxes (Pteropus poliocephalus and P. scapulatus)

Adult male flying foxes Pteropus poliocephalus and P. scapulatus were captured in south-east Queensland and kept in outdoor enclosures. Testicular size (TS), plasma testosterone concentrations (PTC) and body weight (BW) were measured over 1-year periods. Testicular recrudescence in P. poliocephalus began before the summer solstice and TS was greatest during mid-March (autumn) and lowest from July to September. Large increases in PTC were observed in all individuals approximately 1 month after the peak in TS. BW also increased around the time of the mating season, changes being correlated significantly with changes in TS. Mating occurred between April and June, and births from late October to late November. In P. scapulatus, TS was greatest in the spring (October) and least in the autumn (February to May); PTC fluctuated throughout the year in this species but, unlike P. poliocephalus, did not show a single large increase in the mating season. BW showed a similar seasonal pattern to that seen in P. poliocephalus, being greatest at the time of greatest TS. Mating occurred in October to November, and births in autumn. In captivity, in outdoor enclosures, these species maintained the seasonal reproductive patterns observed in the wild. The 2 species respond differently to the same environmental cues in terms of regulation of the timing of their breeding seasons.     

Empirical evidence of a trade-off between reproductive effort and expectation of future reproduction in female three-spined sticklebacks

Optimal life-history models generally predict that the reproductive effort of iteroparous organisms may increase with age, as their expectation of future reproduction decreases. The population of three-spined sticklebacks (Gasterosteus aculeatus) in the Camargue (Rhone River Delta, France) is annual, all adults dying after their first breeding season. As the three-spined stickleback is a multiple spawner, we tested the hypothesis that reproductive effort may increase during the breeding season on field data. From 1987 to 1998, 653 female sticklebacks were collected in the field during the breeding seasons. The body size, body weight and weights of the liver, gonads and carcass were measured for these individuals. Only gravid females with mature eggs (176 fish) were included in the analysis. Considering the female three-spined stickleback as a capital breeder, the energetic resources available for allocation between soma and gonads were estimated by its body weight. Somatic condition decreased during the breeding season and reproductive effort (gonad weight relative to body weight) increased. These patterns did not vary significantly between years. These observed variations in reproductive effort during the breeding season can be interpreted as empirical evidence of a trade-off between reproductive effort and expectation of future reproduction.     

Seasonality, body condition, and timing of reproduction in Propithecus verreauxi verreauxi in the Kirindy Forest

Mammals that live in seasonal environments may adjust their reproductive cycles to cope with fluctuations in food availability. Because lemurs in Madagascar experience highly seasonal variation in food availability, we examined the effects of fluctuating food availability on body condition and reproduction in one of the larger living species, Verreaux's sifaka (Propithecus verreauxi verreauxi), in the Kirindy Forest of western Madagascar. Seven years of demographic data were combined with an intensive study of 25 individuals over the course of 18 months. In contrast to other populations of Verreaux's sifaka, females were found to have greater body mass than males. Both male and female sifaka exhibited significant losses of body mass and fat during the dry season. Females were more likely to give birth and successfully wean an infant when they had higher body mass during the mating season. They mated during the periods of high and declining food availability, gave birth during the lean season, and then timed mid/late lactation with the period of increasing food availability. Thus, we conclude that sifaka follow the "classic" reproductive strategy (sensu van Schaik and van Noordwijk [Journal of Zoology (London) 206:533-549, 1985]).     

Niche separation in Varecia variegata rubra and Eulemur fulvus albifrons: II. Intraspecific patterns

Based on a year-long field study in northeastern Madagascar, I summarize annual patterns of niche use (food patch size, diet, forest height, and forest site) in two sympatric lemurs, Varecia variegata rubra and Eulemur fulvus albifrons. Furthermore, I examine intraspecific patterns of niche use according to sex, season, and reproductive stage in these two lemurs that differ in terms of energetic investment in reproduction. Lemurs as a group provide a special opportunity to test hypotheses concerning sex differences in niche use. Due to their body size monomorphism and seasonal, synchronous pattern of breeding, it is possible to directly evaluate whether sex differences in diet reflect high energetic investment in reproduction by females. Results confirm the hypothesis that intraspecific variation in niche use (e.g., sex differences, seasonal differences) would be more pronounced in V. v. rubra than in E. f. albifrons, due in large measure to the former's relatively high energetic investment in reproduction: 1a) Dietary sex differences in V. v. rubra are most pronounced during costly reproductive stages and involve acquisition of low-fiber, high-protein plant foods. Females of both species consume more seasonally available low-fiber protein (young leaves, flowers) relative to conspecific males during the hot dry season, but only in V. v. rubra females is this pattern also evident during gestation and lactation. 1b) The diets of female V. v. rubra and female E. f. albifrons are more similar to each other than are the diets of conspecific males and females in the case of V. v. rubra. This is not uniformly the case for female E. f. albifrons. This finding confirms a hypothesis put forward in Vasey ([2000] Am J Phys Anthropol 112:411-431) that energetic requirements of reproductive females drive niche separation more than do the energetic requirements of males. 1c) Both species synchronize most or all of lactation with seasonal food abundance and diversity. E. f. albifrons shows a more protracted period of synchrony, and this may contribute to its wide biogeographic distribution in Madagascar. 2) Sex differences and seasonal differences in microhabitat use reflect intraspecific patterns of thermoregulation, predator avoidance, and in the case of V. v. rubra, reproduction. One important factor selecting for body size monomorphism in lemurs appears to be the tight synchrony between lactation and periods of food abundance afforded by annual, seasonal breeding.     

Collection, seminal characteristics and chilled storage of spermatozoa from three species of free-range flying fox (Pteropus spp.)

This study reports observations on the collection and characteristics of semen from free-range populations of flying fox in Brisbane, Australia. Semen was successfully recovered by electroejaculation from 107 of 115 wild flying foxes (Pteropus alecto, Pteropus poliocephalus and Pteropus scapulatus). A proportion of ejaculates collected from all three species contained seminal vesicle secretions, the incidence of which appeared related to breeding season. Ejaculate volume was small (5--160 microL), requiring a specialised collection vessel and immediate extension to avoid desiccation. Sperm morphological abnormalities and characteristics are described for the first time. In two species (P. scapulatus and P. alecto), sperm quality varied with breeding season. Dilution in Tris-citrate-fructose buffer and subsequent incubation (37 degrees C) of Pteropus semen for 2-3h appeared to have a negative impact on sperm motility and the percentage of sperm with intact plasma membranes and acrosomes and represents a concern for the potential development and use of assisted breeding technology in these species. Preliminary attempts to develop a short-term chilled preservation protocol for flying fox semen revealed that sperm viability (percentage motility and percentage live sperm with intact acrosomes) was significantly reduced after 102 h chilled storage at 5 degrees C; nevertheless, approximately 40% of the spermatozoa were still motile and contained intact acrosomes. Glycerol was neither protective nor detrimental to sperm survival during chilled storage. Microbial flora of the prepuce, urethra and semen of all species were isolated and their antibiotic susceptibility tested. Tetracycline, penicillin, ciprofloxacin, and ceftazidime were the most effective antibiotics in preventing growth of all identified bacteria; however, their effects on sperm survival were not investigated.     

Indirect Evidence that Flying Foxes Track Food Resources Among Islands in a Pacific Archipelago

Although flying foxes (fruit bats in the genus Pteropus ) in continental forests often fly between scattered resources, little is known about their ranging behavior among islands. The inhospitable water matrix that surrounds the food patches (islands) in archipelagos may prevent flying foxes from tracking resources as efficiently as their counterparts on larger landmasses do. Our aim in this study was to determine whether the abundance of foraging flying foxes ( Pteropus tonganus ) reflected food availability on islands in the Vava'u archipelago of Tonga, regardless of island size and isolation. Overall, food availability was the strongest determinant of flying fox abundance, and spatial aspects of the islands (land area within 10 km) had only a small influence. Food availability appears to regulate flying fox abundance only when food resources are low, but when food sources are plentiful, flying fox abundance may be high or low. These results provide indirect evidence that flying foxes are able to track food resources efficiently in an archipelago, and the water matrix that surrounds the food patches (islands) is not a strong deterrent for foraging animals.     

Female red squirrels fit Williams' hypothesis of increasing reproductive effort with increasing age

Williams predicted that reproductive effort should increase as individuals age and their reproductive value declines. This simple prediction has proven difficult to test because conventional measures of energy expenditure on reproduction may not be a true reflection of reproductive effort. We investigated age-specific variation in female reproductive effort in a stable population of North American red squirrels where energy expenditure on reproduction is likely to reflect actual reproductive effort. We used seven measures of reproductive effort spanning conception to offspring weaning. We found that females completed growth by age 3 and that reproductive value decreased after this age likely because of reproductive and survival senescence. We therefore, predicted that reproductive effort would increase from age 3 onwards. The probability of breeding, litter mass at weaning, and likelihood of territory bequeathal were all lower for 1- and 2-year-old females than for females older than 3 years, the age at which growth is completed. That growing females are faced with additional energetic requirements might account for their lower allocation to reproduction as compared with older females. The probability of attempting a second reproduction within the same breeding season and the propensity to bequeath the territory to juveniles increased from 3 years of age onwards, indicating an increase in reproductive effort with age. We think this increase in reproductive effort is an adaptive response of females to declining reproductive values when ageing, thereby supporting Williams' prediction.     

Father–offspring phenotypic correlations suggest intralocus sexual conflict for a fitness-linked trait in a wild sexually dimorphic mammal

In sexually dimorphic and polygynous mammals, sexual selection often favours large males with well-developed weaponry, as these secondary sexual characters confer advantages in intrasexual competition and are often preferred by females. Little is known, however, about the effects of sexually selected paternal traits on offspring phenotype in wild mammals, especially when considering that shared phenotypic traits and selection can also differ greatly between genders. Here, we conducted molecular parentage analyses in a long-term study population of mountain goats (Oreamnos americanus), an ungulate exhibiting high sexual dimorphism in mass, to first assess the determinants of yearly reproductive success (YRS) in males. We then examined the effects of paternal characteristics on offspring mass at 1 year of age. Paternity was highly skewed, with 9 per cent of 57 males siring 51 per cent of 96 offspring assigned over 12 years. Male YRS increased with age until apparent reproductive senescence at 9 years, but mass was a stronger determinant of siring success than age, horn length or social rank. Mass of sons increased with paternal mass, but the mass of daughters was negatively related to that of their father, a finding consistent with recent theory on intralocus sexual conflict. Because early differences in mass persisted to early adulthood, sex-specific effects of paternal mass can have important fitness consequences, as adult mass is positively linked with reproduction in both sexes. Divergent father–offspring phenotypic correlations may partly explain the maintenance of sexual dimorphism in mountain goats and the large variance observed for this homologous trait within each gender in polygynous mammals.     

Reproductive constraints,not environmental conditions,shape the ontogeny of sex‐specific mass–size allometry in roe deer

In polygynous mammals, sex‐specific patterns of body growth are linked to divergent selection pressures on male and female body size, resulting in sexual dimorphism (SD). For males, reproductive success is generally linked to body size, hence, males should prioritise early growth. For females, reproductive success is linked to resource availability, so they may adopt a more conservative growth tactic. Using longitudinal monitoring of known‐age animals in two contrasting populations and an allometric approach to disentangle the relative contribution of structural size and physiological condition to SD, we addressed these issues in the weakly polygynous roe deer. Despite very different environmental conditions, we found remarkably similar patterns in the two populations in the mass–size allometric relationship at each life history stage, suggesting that relative allocation to structural size and physiological condition is highly constrained. SD in structural size (indexed by hind foot length) involved sex‐specific growth trajectories governed by a single mass–size allometric relationship during the juvenile stage, such that males were both bigger and heavier than females. In contrast, SD in physiological condition (indexed by the allometric relationship between body mass and hind foot length, expressed as body mass for a given body size) developed markedly during the sub‐adult stage in relation to sex differences in the timing of first reproduction. Among adults, males were heavier for a given size than females, suggesting that, relative to females, males express a capital breeder tactic, accumulating fat reserves to offset reproductive costs. By the senescent stage, SD in physiological condition had disappeared, with both sexes governed by a single allometric relationship, suggesting more rapid senescence in males than females. Individuals born into poor cohorts were generally lighter for a given size, indicating growth priority for skeletal size over physiological condition in both sexes. However, sex differences in cohort effects among sub‐adults resulted in lower size‐specific SD in poor cohorts, indicating that body condition of sub‐adult females is buffered against environmental harshness. We conclude that sex‐differences in reproductive tactics impose constraints on the ontogeny of SD in roe deer, leading to sex‐specific trajectories in structural size and physiological condition.     

Reproductive energetics of the nectar-feeding bat<Emphasis Type="Italic"> Glossophaga soricina</Emphasis> (Phyllostomidae)

Pregnancy and lactation are energetically demanding periods for female mammals. Unique amongst mammals, bats have to allocate considerable amounts of energy into their offspring because juveniles cannot be weaned until they are capable of flying at almost adult size. Similar to other bat species, female nectar-feeding bats should increase their energy intake after parturition to meet the energy demands of offspring growth. However, previous studies have shown that nectar-feeding bats differ from other similar-sized bats in having a much higher metabolic rate. Therefore, I examined how nectarivorous bats respond to the energetic challenge of reproduction. In this study, the daily energy intake of pregnant and lactating Glossophaga soricina was measured during a 6-week period prior to and a 10-week period after parturition. Body mass of G. soricina increased linearly until parturition. Within the same time period, daily flight time decreased and daily energy intake remained constant. Probably, the reduced flight activity of pregnant bats compensated for the increased power requirements of flight, thus resulting in an almost constant daily energy turnover. During 35 days after parturition, neither flight time, body mass nor daily energy intake of lactating females changed significantly. On average, the daily energy intake of pregnant, lactating or non-reproducing G. soricina was not significantly different. Possibly, for unknown reasons, female G. soricina maintain a daily energy intake of a constant high level during and beyond reproduction.     

Intersexual differences in energy expenditure of Anolis carolinensis lizards during breeding and postbreeding seasons

Although the amount of energy that males and females invest in reproduction is an integral component of theories explaining the evolution of particular mating strategies, few studies have actually determined the amount of energy that each sex allocates to reproduction. We compared how energy is expended by male and female Anolis carolinensis lizards during both the breeding and postbreeding seasons. We used laboratory respirometry to determine resting metabolic rates (RMRs) of inactive, freshly captured lizards and the doubly labeled water technique to determine field metabolic rates (FMRs) of free-ranging lizards. Both RMRs and FMRs were influenced by body mass but not by sex. Season did not influence FMRs; however, RMRs of both sexes increased approximately 40% from the breeding to the postbreeding season. The seasonal increase in RMRs was attributed to a postreproductive increase in feeding rate and specific dynamic action. We used RMRs, FMRs, and thermal profiles of lizards to calculate energy budgets for breeding and postbreeding seasons. Energy budgets partitioned daily field energy (DFE; calculated from FMRs) into daily activity energy (DAE) and daily resting energy (DRE; calculated from RMRs). Energy expended for reproduction was estimated as DAE during the breeding season plus egg production (for females). Despite males having 40% greater body mass, females expended 46% more energy for reproduction than did males (906 and 619 J/d, respectively). Total metabolizable energy (TME=DFE+egg production for females) expended during the breeding season was similar for males and females (1,280 and 1,365 J/d, respectively). Although TME of females decreased 44% from the breeding to the postbreeding season (1,365 vs. 766 J/d), TME of males was similar during both seasons (1,280 vs. 1,245 J/d). There were both seasonal and sexual differences in DRE and DAE. Compared with most lizards from semiarid/desert habitats, A. carolinensis in a temperate habitat expends more total energy during the breeding season, allocates more energy to eggs, and appears to have more total energy available for reproduction.