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Circulating leptin levels are elevated during the later stages of pregnancy in mammals, suggesting that maternal leptin may play a role in maintenance of pregnancy and/or preparation for parturition and lactation. The regulation and source of circulating leptin during pregnancy remains undetermined, but leptin mRNA levels increase in adipose tissue during this time in some species. Considerable controversy exists whether placenta is also a leptin-secreting tissue during pregnancy. Here, we directly demonstrate that leptin secretion rates from mouse adipose tissue in vitro are decreased during early pregnancy and up-regulated during late pregnancy and lactation. Changes in leptin secretion rates in vitro paralleled those of circulating leptin in vivo during gestation. Subcutaneous implants of estradiol or corticosterone into lactating mice for 48 h stimulated adipose leptin secretion rates in vitro to the level of that in pregnant mice. However, corticosterone, but not estradiol, increased leptin secretion when added to isolated adipose tissue in vitro. Placentae obtained at two stages of pregnancy did not secrete leptin in vitro, either when acutely isolated or when dissociated into cells for long-term cultures. Placental tissue (or cells) secreted progesterone, however, demonstrating placental viability. We conclude that hyperleptinemia during late pregnancy in mice primarily results from corticosterone-dependent up-regulation of leptin secretion from adipose tissue, and that the placenta does not contribute to leptin secretion. The initial decrease in leptin secretory rates from adipose tissue during early pregnancy may facilitate energy storage for the subsequent, increased metabolic demands of later pregnancy and lactation.

The social structure of an Old World tent-making bat Cynopterus sphinx (Megachiroptera), was investigated in western India. A combination of census and mark–recapture data over 2 years (1996–98) was used to infer the form of the mating system, compositional stability of social groups and mode of new social group formation. The breeding population of C. sphinx was subdivided into diurnal roosting colonies, each of which contained one to five discrete roosting groups and often one or more solitary bats in adjacent roosts. Bats most frequently roosted in stem tents constructed in the flower/fruit clusters of the kitul palm Caryota urens. Temporal variation in social structure was assessed using visual census data for a subset of the study population over 3 years (1995–98) spanning six consecutive reproductive periods. The sex and age composition of diurnal roosting groups indicated a polygynous harem-forming mode of social organization, as groups invariably contained a single adult male, 1–37 reproductive females and their dependent young (n = 33 harems). Harem size averaged 6.1 adults in the wet season (n = 19, SD = 3.5) and 13.6 adults in the dry season (n = 14, SD> = 8.5). The same harem social configuration was maintained year-round, despite a high degree of synchrony and seasonality in the timing of reproduction. Juveniles of both sexes dispersed after weaning and sexually immature bats were never present in harems at the time of parturition. Adult females often remained associated as roostmates from one parturition period to the next, and group cohesion was unaffected by turnover of harem males. Adult females frequently transferred among roosts within the same colony, and harems underwent periodic fissions and fusions. The founding of new harems most often resulted from the fissioning of previously cohesive harems within the same colony. However, some harems contained disproportionate numbers of yearling females, indicating that new groups are also founded by nulliparous females of the same age cohort. A significant degree of heterogeneity in age composition among harems was revealed in the 1998 dry season, but was unrelated to age-stratification of tent roosts. Although formation of new harems may be non-random with respect to age composition of the founders, founding events are not restricted to newly created tents and often involve recolonization of previously occupied roosts.

We evaluated organic and macromineral composition of selected fruits and leaves consumed by the short-nosed fruit bat, Cynopterus sphinx in South India. Results of principal components analysis (PCA) comparing soluble carbohydrates, crude protein, and crude fats indicate a higher percentage of protein in leaves and a higher percentage of carbohydrates and lipids in fruits. However, results of a paired t test comparing these organic components indicated no differences between fruits and leaves. Among the fruits analyzed, Musa x paradisiaca had the highest percentage of carbohydrates, and Psidium guajava had the highest percentage of lipids. Leaves of Cassia fistula, Moringa oleifera, coccinia cordifolia, and F. religiosa had the highest percentage of protein. PCA of selected macrominerals (Ca, Na, K, and P) indicate higher levels of Ca in leaves than in fruits. Results of t tests comparing these macrominerals revealed a difference between fruits and leaves for Ca, but not for the other macrominerals. Among the leaves analyzed, Ca was highest in mature leaves of C. fistula and lowest in leaves of F. religiosa. Leaves of M. oleifera and fruits of Achras sapota were highest in sodium. Among fruits analyzed for macrominerals, Ca was highest in F. bengalensis and lowest in Prosopis juliflora, A. sapota, and M. x paradisiaca. Potassium levels were highest in leaves of C. cordifolia and fruit pods of Prosopis juliflora. Phosphorus content was highest in leaves of M. oleifera and fruits of M. x paradisiaca. The relatively high concentrations of protein and calcium in leaves eaten by C. sphinx supports the hypothesis that leaves are important dietary sources for this plant-visiting bat.

Kerivoula pellucida is a small (4.5 g) vespertilionid bat, which forages in the understorey of tropical rainforests in Southeast Asia. Most echolocation signals are characteristic for the subfamily (very high-frequency, broadband FM calls (sweeping down from 178 kHz to 58 kHz) of low intensity and short duration (< 3.0 ms)), but this species also produces stereotypical calls of a markedly different structure. The arrangement of signal elements in these calls is unusual, and we present here a detailed analysis of calls from thirteen adult individuals (nine males and four females). These multiharmonic calls are of much lower frequency (fundamental peak frequency of 36 kHz), higher intensity and longer duration (12.0 ms) than the orientation calls, and are composites comprising a long quasi-constant frequency (QCF) syllable followed by an upward frequency modulated sweep (UFM). From considerations of signal design and preliminary field observations, we consider possible functions of these calls.

Body composition of little brown bats, Myotis lucifugus was quantified to evaluate the proximate costs of reproduction in a small temperate mammal. Changes in the major components of body composition (water, lean dry, and fat mass) during the breeding season were analyzed in adult females and in young during the postnatal growth period. Changes in body composition of reproductive females were small relative to their level of reproductive effort, suggesting that most of the increased costs of reproduction are not met by mobilizing body reserves. The reduction of lean dry and fat mass in adult females suggest that mineral and fat stores are mobilized to promote high postnatal growth rates in pups. Changes in the gastrointestinal (GI) tract of adult females support the hypothesis that increased food intake and assimilation are the primary morphological and physiological responses to the increased energy demands of lactation. A two-week period of linear growth during the postnatal period when pups wer...

Patterns of dispersion and site fidelity were investigated in a tent-roosting population of the short-nosed fruit bat, Cynopterus sphinx (Megachiroptera), in southern India. A local population of C. sphinx occupied diurnal roosts in a variable subset of 45 stem tents constructed within the dense foliage of mast trees (Polyalthia longifolia). Individually marked tent-roosting bats were visually censused over the course of a 38-d interval spanning the postpartum oestrus period. On any given day, 33.3–85.7% (mean = 60.8%, SD = 14.2) of adult males roosted singly, with the remainder holding harems of 1–10 breeding females (mean = 3.01, SD = 0.79). Average harem sex ratio was 2.8-fold higher than the adult sex ratio of the total tent-roosting population within the study area, indicating the potential for a high variance in male mating success within a single breeding season. Bats of both sexes typically occupied one primary tent, interspersed with shorter periods of residency in alternate tents. Males exhibited a significantly higher degree of roost fidelity than females. Some females roosted sequentially with different males and with different combinations of females, whereas others remained continuously associated with a single male and/or particular female roostmates over the duration of the census period. There were no statistically significant relationships between physical characteristics of tents and rates of occupancy by males or females. Intermittent transfers by females between groups suggest that the defence of diurnal roosts by males represents a more profitable mating strategy than the direct defence of compositionally labile female groups.

Echolocation signal design in nine syntopic vespertilionid bats from the Malaysian rain forest understorey was studied. Four species of Kerivoula, two species of Phoniscus (Kerivoulinae) and three species of Murina (Murininae) all emitted calls that are typical of species that glean insects from surfaces: broadband, frequency-modulated (FM) calls of low intensity and short duration. However, calls were highly distinctive in the use of very large bandwidths (range: 89–123 kHz) and extremely high frequencies (start frequency 152–180 kHz; end frequency 43–86 kHz). Furthermore, calls were produced in groups of 2–15 at very high pulse repetition rates (37–105 Hz). The functional significance of these characteristics with respect to foraging strategy is discussed. Large signal bandwidths facilitate highly accurate target localization in terms of both range and angle estimation and can thus be interpreted as an adaptation to foraging in the highly cluttered environment of the forest understorey. The use of high frequencies so far in excess of those seen in other FM gleaning bats of the same size is less easily explained, but may represent a mechanism by which these species distinguish prey items using echolocation alone, without recourse to visual or auditory cues. Species exhibited differences in echolocation parameters, particularly the end frequency and the number of calls per group of calls. The two subfamilies differed from each other in multivariate space derived from echolocation parameters; calls of the Murininae were of lower frequency than the Kerivoulinae and were typically produced in smaller groups or singly. Within the subfamilies there was considerable overlap between species of Murina, but the Kerivoula spp. were clearly distinct from one another but not from the two Phoniscus spp.

The short-nosed fruit bat, Cynopterus sphinx, begins to visit fruit-bearing trees about 30 min after sunset. Individual bats often hover near or land on fruits or on nearby branches to remove whole or parts of fruits with their mouth. These bats seldom remain in the fruit-bearing trees to feed, but instead carry fruits to feeding roosts, repeating this behaviour several times throughout the night. Analysis of the temporal distribution of feeding behaviour has revealed two peaks of activity, one in the pre-midnight hours when bats fed mostly on ‘steady state’ fruits, and another during the post-midnight hours when bats fed on ‘big-bang’ fruits. Only solitary bats visited and fed on species with steady state fruiting phenologies, whereas groups of bats regularly visited and fed on species with big-bang fruiting phenologies. Thus, plant species which produce large numbers of fruits appear to promote group foraging during the latter hours of the night. It is suggested that the temporal use of available fruits in south India made it possible for C. sphinx to successfully exploit them, and thereby reduced interference competition with conspecifics.

We evaluated seasonal dietary variation of the Indian pygmy bat, Pipistrellus mimus , from southern India, by collecting biweekly fecal samples from beneath a maternity roost for 1 year and comparing these samples with insects collected using a light trap on the same dates. Our analyses indicate that P. mimus fed on coleopterans (26.3% volume), dipterans (20.4%), homopterans (17.5%), lepidopterans (13.6%), formicids (10.5%), hemipterans (5.7%), and isopterans (5.1%) and thus are generalized feeders. One unidentified scarabaeid species made up much of the coleopteran material and was abundant in the diet of P. mimus from late April through late August. Formicid alates were consumed throughout the year but were especially important from late July through September. Isopteran alates were important in September and October. Relative proportions of insects that were eaten differed significantly from insects captured in a light trap. In other studies, investigators interpreted such differences as evidence of selection; however, we argue that most insect traps do not provide reliable data on availability and that bats may feed in other areas than where the insects are captured. Our results indicate that >20 fecal pellets are needed in any one sample to determine important dietary items eaten by this species, although this sampling protocol likely overlooks minor food items.

To investigate constraints on males associated with maintenance of harems, we used a time-energy approach to explore the relationship between temporal occupancy of roosts and nightly foraging activity in male and female greater spear-nosed bats, Phyllostomus hastatus. Harem males and females differed significantly in frequency and duration of nightly foraging bouts. Harem males spent less time away from their roost at night and departed and returned more frequently than did other members of the harem. Adult females and one subadult male foraged mostly during the first 2-3 h after sunset. Total time spent foraging by harem males, adult females, and a subadult male, averaged 98, 152, and 115 min per night, respectively. On average, harem males engaged in 7.2 randomly spaced foraging bouts per night, compared with 2.1 bouts for harem females and 2.0 bouts for a subadult male. Field metabolic rate, expressed on a mass-specific basis (ml CO2 g-1 h-1), averaged 3.4 ± 0.69 SD for harem males, 3.0 ± 0.85 for adult females, 1.9 ± 0.08 for a subadult male. These values varied predictably but did not differ significantly. Similarly, daily energetic expenditure, expressed on a whole-animal basis (kJ/day), averaged 168.6 ± 32.5 for harem males and 124.9 ± 38.9 for adult females but did not differ significantly. Because harem males allocate less time to nightly foraging activity compared with adult females, males should have more time and energy available for vigilance at the roost and/or the defense of females.

Information on the feeding habits of the the lesser dog-faced friut bat, Cynopterus brachyotis, was obtained by the collection of food remains directly beneath daytime and feeding roosts. The bats were found to feed on the fruits of 54 plant species, the leaves of 14 species and the flower parts of four species. The seasonal phenological differences among congeneric plant species led to steady production of fruit throughout the year and the data suggets that Ficus spp. are a key component in the diet. Judging from its wide selection of fruits, C. brachyotis, is considered to be an important seed disperser. Folivory in C. brachyotis appears to be more common than previously thought. Of the leaves consumed by the bats, seven species belonged to the family Leguminosae, followed by Myrtaccae, Moraccae, Rhizophoacae and Euphorbiaccae. Friuts in general, provide an energy rich diet for phytophagous bats but most are low in protein. In contrast, leaves consumed by bats have a relatively high protein content. We suggest that folivory (by leaf fractionation) should be energetically more advantageous than the ingestion of large amounts of low protein friut or the active pursuit of mobile insects.

We describe the size and developmental state at birth, examine factors which influence intra- and inter-year variation in post-natal growth, and derive age-predictive equations and logistic growth parameters for the eastern pipistrelle bat, Pipistrellus subflavus. Ambient and roost temperature and insect abundance were lowest, and precipitation highest, before and during the post-natal growth period in 1982, as compared to 1981. We found no sex differences in the size of pups at birth or in post-natal growth rates in a given year; however, body mass and length of forearm of neonates differed significantly between years, being smaller in 1982 than in 1981. During the first two weeks of post-natal growth in both years, the body mass and length of forearm of pups increased linearly at mean rates of 0.15 g.day−1 and 1.12 mm.day−1, respectively. Post-natal growth rates for length of forearm and body mass were the same during the early linear phase of growth, but the intercept for body mass was less and growth trajectories were more variable in 1982. Length of forearm was the least variable and thus most reliable character for estimating age of P. subflavus pups during the first 14 days of post-natal growth. When length of forearm was used in combination with mean changes in the length of the fourth metacarpal-phalangeal epiphyseal gap, we derived reliable age-estimation equations ranging from 1 to 45 days. Litter size averaged 1.96 and 1.93 at birth, but this was reduced to one by the time pups became volant. At birth, litter mass represented 44, and 54% of the post-partum mass of females in 1981 and 1982, respectively. This maternal effort during pregnancy is among the highest reported for bats. At weaning, the body mass of pups approached 80% of adult post-partum body mass, and the length of forearm exceeded 90% of adult female size. Our results suggest that size at birth and post-natal growth are influenced by ambient and roost temperature, rainfall, and the quantity of insects available to pregnant and lactating females. Thus, energetic constraints on both mothers and pups can lead to inter-year and inter-individual differences in maternal effort and post-natal growth of pups.

Changes in body mass, fat mass, lean dry mass, and energy content of little brown bats, Myotis lucifugus (LeConte), captured at a cave in southern Vermont, were quantified during the pre-hibernatio...

We measured postnatal changes in wing morphology and flight capability in free-ranging Phyllostomus hastatus (Chiroptera: Phyllostomidae). Area of wing, wingspan, and aspect ratio increased until volancy, while wing loading reached its lowest values at and after weaning. Young began to fly in their 7th week, when wing area, aspect ratio, and wingspan had reached 84, 95, and 89%, respectively, of values for adult females. Foraging time increased as young achieved volancy. Estimates of flight speed and power requirements from theoretical predictions yield flight speeds that are lowest around weaning, power requirements that increase consistently, and a cost of transport that decreases to a constant level at weaning. Thus, wing morphology of young may function to maximize maneuverability and minimize energy requirements during the vulnerable period of first flight. Wing loading of adult females fluctuated greatly with seasonal changes in body mass. Wing loading of nonreproductive females, a factor of their greater wing area and lower body mass, was significantly lower than in males. An upper limit on the ability to accomodate increased mass may affect wing morphology in this species.

This study examined the tent-making behaviour and social organization of the lesser dog-faced fruit bat, Cynopterus brachyotis (Muller), in Peninsular Malaysia. Cynopterus brachyotis alters the leaves of eight species of broad-leafed palms (Palmae) to form palmate umbrella tents, the leaves of Scindapsus aureus Engl. (Araceae) to form apical tents, and the leaves of Dracaena fragrans Ker-Gawl (Liliaceae) to form conical tents. Cynopterus brachyotis also severed the roots of Cymbidium finlaysonianum Lindl (Orchidaceae), and the leaves and stems of Philodendron lacerum Schott (Araceae) to form stem tents. In addition, C. brachyotis also roosted in unmodified plants. Cynopterus brachyotis possesses a harem type of social organization, but it was not possible to distinguish between a resource-defence polygyny and female-defence polygyny mating system. Our observations indicate that dominant males construct tents, and recruit females for the purpose of gaining reproductive access.

Growth rates of mammalian young are closely linked to the ability of the mother to provide nutrients; thus, milk composition and yield provide a direct measure of maternal investment during lactation in many mammals. We studied changes in milk composition and output throughout lactation in a free-ranging population of the omnivorous bat, Phyllostomus hastatus. Fat and dry matter of milk increased from 9 to 21% and from 21 to 35% of wet mass, respectively, throughout lactation. Energy increased from 6 to 9 kJ.g-1 wet mass, primarily due to the increase in fat concentration. Total sugar levels decreased slightly but non-significantly. Mean sugar level was 4.0% of wet mass. Protein concentration increased from 6 to 11% of wet mass at peak lactation and then decreased as pups approached weaning age. Total milk energy output until pups began to forage was 3609 kJ. Milk levels of Mg, Fe, Ca, K, and Na averaged 0.55 %/- 0.26, 0.23 +/- 0.2, 8.75 +/- 4.17, 5.42 +/- 2.11, and 9.87 +/- 4.3 mg.g-1 dry matter, respectively. Of the minerals studied, calcium appears to be most limiting in this species. The high degree of variability in foraging time, milk composition and milk yield between individuals at the same stage of lactation could potentially yield high variance in reproductive success among females of this polygynous species.

This study reports developmental changes in the vocal signals produced by wild-caught pre-volant and volant Myotis lucifugus. Audio recordings were made from young animals (1-33 days old) and adults (over 1 year of age). The animals were removed from an attic maternity roost and studied individually in a room below. To stimulateflight-related behaviours, animals were released from a launching platform via a hingedfloor, and their vocalizations were recorded as they approached a soft foam pad below. When the hinged floor opened, the youngest animals (1-4 days old) typically dropped onto the pad, but older animals either flapped their wings to achieve some horizontal displacement (>4 days) or sustained horizontal flight (>17 days). Vocalizations recorded under these conditions showed frequency modulation characteristic of adult echolocation sounds, even in animals as young as 4 days. Ontogenetic trends showed an increase in sound frequency, an increase in sound repetition rate and a decrease in sound duration as the animals matured. These data are discussed in the context of the development of echolocation behaviour in bats. ? 1997 The Association for the Study of Animal Behaviour Insectivorous bats use echolocation to orient in the environment and capture prey. The bat's vocal apparatus functions as a sonar transmitter of ultrasonic signals that reflect well oV small objects, and the bat's auditory system functions as a sonar receiver that determines the angular position and distance of targets (GriYn 1958; Schnitzler & Henson 1980; Moss & Schnitzler 1995). Before young animals begin to hunt insects successfully during their first flights, they must be capable of producing high frequency sonar cries and of processing the spatial information con- tained in the sonar echoes. Thus, the vocal pro- duction system and the auditory system must develop and operate in concert for the bat to use its biological sonar. Data on the vocalizations of young bats exist for several families, including Vespertilionidae

We examined patterns of variation in the mineral content of the wing skeleton of the Mexican free-tailed bat, Tadarida brasiliensis. We ashed humeri, radii, metacarpals II-V, and phalanges of digit III, and quantified mineralization differences among elements at specific ages, and ontogenetically for each element. The most mineralized elements are the humerus and the radius, followed by the metacarpals, of which the third and fifth are the most mineralized. The proximal and middle phalanges of the third digit exhibit the lowest mineral content, and the distal phalanges have no mineral content according to our ashing protocol. Histological examination shows a thin (< 10 μm) shell of unmineralized osteoid surrounding a cartilaginous core in distal phalanges. Mineral content of each bone increases linearly with age during post-natal development, but there are differences in the rate and extent of this increase among the different elements. The mineralizaton differences we observed parallel substantially different bone loading patterns found in different parts of the wing in other studies. The humerus and radius are subjected to large torsional loads during flight, while the metacarpals and phalanges experience dorsoventral bending. The high mineral content of the humerus and radius and the low mineral content of the metacarpals and phalanges may resist torsion proximally and promote bending distally. Furthermore, the decrease in mineral content along the wing's proximodistal axis decreases bone mass disproportionately at the wing tips, where the energetic cost of accelerating and decelerating limb mass is greatest.

We examine how litter mass in bats varies with respect to wing loading, an important aerodynamic aspect of flight. From geometric proportions, litter mass should scale to wing loading by an exponent of three. Conversely, analysis of aerodynamic consequences of carrying extra mass suggests that an exponent significantly less than three would be selectively advantageous. Our results show that Megachiroptera and Microchiroptera differ in the relationship between litter mass and wing loading. Litter mass in megachiropterans scales as expected by geometric proportions, whereas litter mass in microchiropterans, as a group, and for individual families, scales as expected if aerodynamic consequences of flight influence litter mass more than size constraints. Thus, selection pressures on reproductive traits appear to differ between the two suborders of bats.

Daily and seasonal activity budgets of adult female eastern pipistrelle bats, Pipistrellus subflavus, were quantified at a maternity colony in eastern Massachusetts. Ambient and roost temperatures were recorded. Activities of roosting bats were monitored using an infrared-sensitive video camera. Scan sampling was used to analyze video records taken at weekly intervals from mid-May to late July 1989. Analysis of day-roosting indicates that adult females spend an average of 77% of their time at rest, 16% alert, 7% grooming, and

Palmate umbrella tents used by tent-making bats in Trinidad, West Indies, were observed in three species of Neotropical palms, Sabal mauritiiformis, Coccothrinax barbadensis and Mauritia flexuosa. Tents were most common in palm leaves that have supporting petioles angled at 50–70° above the horizontal. The shape and volume of tents is influenced mostly by leaf morphology (leaf width and leaflet length) and age of the tent. Tent-crowns varied from being heart-shaped in S. mauritiiformis, oval or round in C. barbadensis and spade-shaped in M. flexuosa. Leaves in which tents were constructed were most often beneath overhanging vegetation, and were generally free of vegetation below, allowing bats to enter and depart from tents without being impeded by the clutter of adjacent vegetation.Singles and small apparent harem groups of two bat species, Artibeus jamaicensis and Uroderma bilobatum, were captured and observed in tents made from the leaves of S. mauritiiformis and C. barbadensis. No bats were observed in tents constructed in leaves of M. flexuosa. The apparent harem social organization in these and other tent-making bat species suggests that leaves modified into tents may provide critical and defendable resources that promote the evolution of polygyny. This hypothesis is based on the observed patchy distribution of suitable palm trees, the inappropriateness of many palm leaves as potential tents and the resultant architecture provided by palmate umbrella tents. We suggest that tent-making is an adult male behaviour.

Disease can play an important role in structuring species communities because the effects of disease vary among hosts; some species are driven towards extinction, while others suffer relatively little impact. Why disease impacts vary among host species remains poorly understood for most multi-host pathogens, and factors allowing less-susceptible species to persist could be useful in conserving highly affected species. White-nose syndrome (WNS), an emerging fungal disease of bats, has decimated some species while sympatric and closely related species have experienced little effect. We analysed data on infection prevalence, fungal loads and environmental factors to determine how variation in infection among sympatric host species influenced the severity of WNS population impacts. Intense transmission resulted in almost uniformly high prevalence in all species. By contrast, fungal loads varied over 3 orders of magnitude among species, and explained 98% of the variation among species in disease impacts. Fungal loads increased with hibernating roosting temperatures, with bats roosting at warmer temperatures having higher fungal loads and suffering greater WNS impacts. We also found evidence of a threshold fungal load, above which the probability of mortality may increase sharply, and this threshold was similar for multiple species. This study demonstrates how differences in behavioural traits among species—in this case microclimate preferences—that may have been previously adaptive can be deleterious after the introduction of a new pathogen. Management to reduce pathogen loads rather than exposure may be an effective way of reducing disease impact and preventing species extinctions. This article is part of the themed issue ‘Tackling emerging fungal threats to animal health, food security and ecosystem resilience’.

Stomach content analysis of 20 pregnant (x body mass=13.4 g) and 18 lactating (x body mass=11.5 g) female Tadarida brasiliensis revealed that the diet, expressed as percent volume, consists largely of lepidopterans, coleopterans, hymenopterans, and dipterans, in decreasing order of importance. We found no significant difference in the diet of pregnant and lactating females when expressed as percent volume. However, when expressed as percent frequency, proportionately more pregnant females fed on lepidopterans, coleopterans, and dipterans than did lactating bats, and proportionately more lactating females fed on hymenopterans. We found no significant differences in the percentages of water, lean dry mass, fat, and energy density in the stomach contents of pregnant and lactating females. Water in stomach contents averaged 62.7% and fat and lean dry mass averaged 22.2% and 15.2%, respectively (expressed as percentage of wet mass); energy density averaged 31.2 kJ g−1 dry mass. This relatively high energy density of stomach contents, as compared to whole insects, can be attributed to the consumption of insects high in fat (especially flying ants) and the abdomens only of moths and beetles (other body parts being discarded). Estimates of nightly food intake increased markedly from mid- to late pregnancy, stabilized or decreased during late pregnancy, and increased again during early to mid-lactation. Average nightly feeding rate doubled from pregnancy to lactation and increased threefold during the first half of lactation. By adjusting our gravimetric estimates of nightly feeding rate upwards by 40% (based on estimates of field metabolic rate), we estimate that the average nightly energy intake of female T. brasiliensis ranges from 57 kJ day−1 in early lactation to 104 kJ day−1 in mid-lactation. These estimates represent nightly feeding rates averaging 39% and 73% of a female's body mass in the period from early to mid-lactation, respectively.

The short-nosed fruit bat (Cynopterus sphinx) creates bell-shaped cavities in flower/fruit clusters of the kitul palm (Caryota urens) by chewing and severing flower and fruit strings. These cavities (stem tents) in which the bats roost are usually about one metre deep and 30 cm in diameter. We observed groups of bats roosting in fully-formed stem tents during the daytime, and the construction and subsequent occupancy of newly formed tent cavities. Stem tents are similar in principle to leaf tents except, instead of being formed when bats chew veins and the surrounding tissues of leaves, stem tents are formed in C. urens when bats completely cut several of the central flower/fruit strings. Flower/fruit strings are mostly severed when they are in an immature stage, at times when they are thin and widely spaced. Once these strings thicken and become heavily-laden with mature fruits, bats cannot penetrate the cluster to sever them. Our observations suggest that a single male enters an immature flower or fruit cluster either from below or the sides and severs the central strings along the peduncle. In early phases of stem-tent construction, C. sphinx severs flower/fruit strings at a rate of about one or two per day, and cluster alteration may continue upwards to two months. Only one immature flower/fruit cluster on a C. urens tree is available for alteration by bats at any given time. That this bat does not roost in the fruit/flower cluster during the day, when a tent is under construction, and the accumulation of chewed flower and fruit strings beneath such a cluster in the morning, suggests that tent construction by C. sphinx is a night-time activity.

Levels of nitrogen, sodium, potassium, magnesium, calcium, and iron were determined for suckling young of known-age, lactating Myotis velifer and Tadarida brasiliensis , and milk from T. brasiliensis nursing known-age young. From birth to weaning, relationships of both nutrient concentrations and nutrient accretion to age are similar for both species but differ for each nutrient and generally reach adult concentration by weaning. While total mass of body sodium and potassium reach adult levels in both species well before weaning, levels of other elements do not. Although comparable data are limited, element concentrations in suckling bats are similar to those for other suckling small mammals and nestling birds. Growth requirements for measured nutrients in these bats are lower than published values for rodents. Most mineral nutrients that were measured in milk for T. brasiliensis varied with stage of lactation and were comparable in concentration with milk of other mammals. Using daily accretion of nutrients and their concentrations in milk, we calculated the minimal mass of milk needed to meet growth requirements in suckling T brasiliensis . Among measured nutrients, calcium is the limiting nutritional factor that determines mass of milk needed by suckling young during postnatal growth.

It has long been accepted that most vertebrate animals meet their vitamin D requirements from exposure of skin to UV-B (UV-B) radiation. Many factors affect this endogenous synthesis of vitamin D, including season, latitude, time of day, age, presence of hair, and degree of skin pigmentation. Most bats roost in dark places by day and forage at night, and thus have little or no potential for sunlight exposure. Notwithstanding, some tropical species are diurnal and are known to roost in the canopy of trees where they may be exposed to sunlight for up to 12 h each day. In this study, two species of captive tropical bats (both species are active at night but one, Rousettus aegyptiacus, roosts in caves, tombs, and buildings, whereas the other, Pteropus hypomelanus, roosts in trees) were evaluated for their ability to endogenously synthesize vitamin D. Following timed periods of sunlight exposure, blood plasma was analyzed using a competitive protein binding assay (CPBA) to determine concentrations of 25-hydroxyvitamin D [25(OH)D], the major circulating vitamin D metabolite. The ability to photoconvert provitamin D (7-dehydrocholesterol, 7-DHC) in the sub-tropical winter was determined using sunlight exposed borosilicate samples of 7-DHC in hourly increments. Finally, both species were evaluated in their preference for a roost site by the release of individuals into sunlight or shade in timed trials. Our results support the hypotheses: (1) when exposed to natural sunlight, both species exhibited an ability to endogenously synthesize vitamin D, although significant differences were found between the two, (2) photoconversion of 7-DHC to previtamin D3 is possible during the mid-day hours of a sub-tropical winter day and (3) captive, cave roosting R. aegyptiacus will choose shaded roost sites while captive P. hypomelanus will show no preference for either shade or sun.

White-nose syndrome (WNS) is an emerging infectious disease devastating hibernating North American bat populations that is caused by the psychrophilic fungus Geomyces destructans. Previous histopathological analysis demonstrated little evidence of inflammatory responses in infected bats, however few studies have compared other aspects of immune function between WNS-affected and unaffected bats. We collected bats from confirmed WNS-affected and unaffected sites during the winter of 2008–2009 and compared estimates of their circulating levels of total leukocytes, total immunoglobulins, cytokines and total antioxidants. Bats from affected and unaffected sites did not differ in their total circulating immunoglobulin levels, but significantly higher leukocyte counts were observed in bats from affected sites and particularly in affected bats with elevated body temperatures (above 20°C). Bats from WNS-affected sites exhibited significantly lower antioxidant activity and levels of interleukin-4 (IL-4), a cytokine that induces T cell differentiation. Within affected sites only, bats exhibiting visible fungal infections had significantly lower antioxidant activity and levels of IL-4 compared to bats without visible fungal infections. Overall, bats hibernating in WNS-affected sites showed immunological changes that may be evident of attempted defense against G. destructans. Observed changes, specifically elevated circulating leukocytes, may also be related to the documented changes in thermoregulatory behaviors of affected bats (i.e. increased frequencies in arousal from torpor). Alterations in immune function may reflect expensive energetic costs associated with these processes and intrinsic qualities of the immunocapability of hibernating bats to clear fungal infections. Additionally, lowered antioxidant activity indicates a possible imbalance in the pro- versus antioxidant system, may reflect oxidative tissue damage, and should be investigated as a contributor to WNS-associated morbidity and mortality.

Disease has caused striking declines in wildlife and threatens numerous species with extinction. Theory suggests that the ecology and density-dependence of transmission dynamics can determine the probability of disease-caused extinction, but few empirical studies have simultaneously examined multiple factors influencing disease impact. We show, in hibernating bats infected with Geomyces destructans, that impacts of disease on solitary species were lower in smaller populations, whereas in socially gregarious species declines were equally severe in populations spanning four orders of magnitude. However, as these gregarious species declined, we observed decreases in social group size that reduced the likelihood of extinction. In addition, disease impacts in these species increased with humidity and temperature such that the coldest and driest roosts provided initial refuge from disease. These results expand our theoretical framework and provide an empirical basis for determining which host species are likely to be driven extinct while management action is still possible.

Solar radiation affects all aspects of the Earth’s environment both directly and indirectly. Radiation from the sun produces a wide range of wavelengths that reach Earth and are absorbed or reflected from animate and inanimate surfaces. Visual, ultraviolet and infrared wavelengths, and solar intensity are the primary variables measured to evaluate their effects on the behavioral, thermoregulatory, and cellular responses of organisms. This review will focus on the effects of solar radiation on animal thermoregulation and the various methods used by scientists to assess the effects of infrared radiation on skin temperatures.

White-nose syndrome (WNS) is an emerging infectious disease affecting several species of hibernating bats in North America and threatening at least one species with regional extinction. The syndrome is associated with a cutaneous fungal infection caused by the newly described fungus, Geomyces destructans (Gd) , reduced fat reserves, atypical behavior, severe wing damage, and high mortality. Six North American bat species are currently affected by WNS, but Gd has been reported from three other hibernating species. Given the rapid geographic spread of Gd , several other species are at risk of infection. G. destructans has also been isolated from nine European bat species. However, no other symptoms of WNS or associated mass mortality of bats associated with this fungus has been observed in Europe. The postulated pathogenic role of Gd is not fully understood, although it appears to disrupt the physiology and alter the behavior of hibernating bats. Given the unique life-history traits of hibernating bats, decimated populations will recover very slowly, if at all, unless the syndrome can be effectively managed. Without implementation of effective management practices ( e.g. , mitigation and recovery), lasting ecological and economic impacts at both regional and national scales can be expected.

Interest in forecasting impacts of climate change have heightened attention in recent decades to how animals respond to variation in climate and weather patterns. One difficulty in determining animal response to climate variation is lack of long-term datasets that record animal behaviors over decadal scales. We used radar observations from the national NEXRAD network of Doppler weather radars to measure how group behavior in a colonially-roosting bat species responded to annual variation in climate and daily variation in weather over the past 11 years. Brazilian free-tailed bats (Tadarida brasiliensis) form dense aggregations in cave roosts in Texas. These bats emerge from caves daily to forage at high altitudes, which makes them detectable with Doppler weather radars. Timing of emergence in bats is often viewed as an adaptive trade-off between emerging early and risking predation or increased competition and emerging late which restricts foraging opportunities. We used timing of emergence from five maternity colonies of Brazilian free-tailed bats in south-central Texas during the peak lactation period (15 June–15 July) to determine whether emergence behavior was associated with summer drought conditions and daily temperatures. Bats emerged significantly earlier during years with extreme drought conditions than during moist years. Bats emerged later on days with high surface temperatures in both dry and moist years, but there was no relationship between surface temperatures and timing of emergence in summers with normal moisture levels. We conclude that emergence behavior is a flexible animal response to climate and weather conditions and may be a useful indicator for monitoring animal response to long-term shifts in climate.

Six bat species in northeastern North America spend winter hibernating in caves/mines. Hibernation is characterized by multi-day periods of torpor, when metabolic rates are reduced. Body fat levels increase from 7 to 27% in the fall, just prior to the onset of hibernation by bats. Some groups of hibernating bats in northeastern North America display abnormal behaviors and increased overwinter mortality called white nose syndrome (WNS). Large numbers of dead bats are found in hibernation sites affected by WNS. Previous research has demonstrated that the increased over-mortality is due to the depletion of depot fat reserves by February, which is in turn caused by greatly shortened torpor bouts. Published studies on hibernating mammals have revealed that moderately high levels of polyunsaturated fatty acids (PUFAs) are required in the diet during the fall for torpor. We thus predicted that bats suffering from WNS have lower levels of PUFAs in their fall diets than those that do not suffer from WNS. We tested this hypothesis by analyzing white adipose tissue (WAT) samples from: (a) little brown bats (Myotis lucifugus) collected from different hibernation sites and (b) big brown bats (Eptesicus fuscus), a species not greatly affected by WNS. Our analyses reveal that M. lucifugus populations prone to WNS have significantly less of the PUFA α-linolenic acid in their fall diets than those where WNS does not occur. The fall diets of E. fuscus contain significantly more of the PUFA linoleic acid than those of M. lucifugus from the same site. Our findings thus support the hypothesis that bats suffering from WNS have lower levels of PUFAs in their fall diets.

We report observations of allomaternal care, or epimeletic (care-giving) behaviour, exhibited by one female toward another, before, during, and after giving birth, in the Rodrigues fruit bat, Pteropus rodricensis (Chiroptera: Pteropodidae). Interactions between the expectant mother and other roost mates, from the onset to completion of birth, were observed continuously over a period of approximately three hours. The attending female assisted the mother when she: (I) intermittently groomed her anovaginal region; (2) grasped her with partially outstretched wings; (3) ‘tutored’ her in a feet-down birthing posture; (4) groomed the emerging pup; and (5) physically assisted the mother by manoeuvring the pup into a suckling position. This is the first report of extensive assistance given by a female bat to another during parturition. We suggest that epimeletic behaviour may be common in the highly social Chiroptera, especially among primiparous or inexperienced females and/or when an expectant mother experiences difficulty during labour and parturition. Our observations support the hypothesis that roosting associations in bats may be as important as relatedness in explaining the occurrence of apparently altruistic acts.

Fifteen species of neotropical and three species of paleotropical bats are known either to roost in or to make tents in over 80 species of vascular plants. We summarize the current knowledge of bat-tent architecture, report two new styles of tents (conical and apical) from the Paleotropics, compare the similarity in tents constructed, or used, by neotropical and paleotropical bats, and consider possible functions of tents. Seven styles of tents are known from the Neotropics, three (conical, palmate umbrella, and apical tents) are known from both the Neo- and the Paleotropics, and one (stem tent) is unique to the Paleotropics. In the Neotropics tent-roosting and/or tent-making appears to be a behavior unique to the diverse microchiropteran family Phyllostomidae (subfamily Phyllostomatinae: tribe Stenodermatini), and in the Paleotropics two members of the megachiropteran family Pteropodidae and one member of the microchiropteran family Vespertilionidae are known to construct or roost in tents. Despite the variety of plant taxa used by bats in tent construction, there appears to be a limited number of different leaf forms that can be altered by bats and used as tents. We suggest that the similarity in tent architecture observed among the neotropical and paleotropical bats is a consequence of convergence in leaf morphology among forest understory plants. The congruence in tent-making/roosting behavior observed in members of the Stenodermatini and the Pteropodidae (genusCynopterus) suggests a phylogenetic influence on these behaviors. The similarity in tent-making and/or tent-roosting behavior and life-history traits (small

The impact of ecology and social organization on genetic structure at landscape spatial scales, where gene dynamics shape evolution as well as determine susceptibility to habitat fragmentation, is poorly understood. Attempts to assess these effects must take into account the potentially confounding effects of history. We used microsatellites to compare genetic structure in seven bat species with contrasting patterns of roosting ecology and social organization, all of which are codistributed in an ancient forest habitat that has been exceptionally buffered from radical habitat shifts. Over one thousand individuals were captured at foraging sites and genotyped at polymorphic microsatellite loci. Analyses of spatially explicit genotype data revealed interspecies differences in the extent of movement and gene flow and genetic structure across continuous intact forest. Highest positive genetic structure was observed in tree-roosting taxa that roost either alone or in small groups. By comparison, a complete absence of genetic autocorrelation was noted in the cave-roosting colonial species across the study area. Our results thus reveal measurable interspecies differences in the natural limits of gene flow in an unmodified habitat, which we attribute to contrasting roosting ecology and social organization. The consequences of ecology and behaviour for gene flow have important implications for conservation. In particular, tree-roosting species characterized by lower vagility and thus gene flow will be disproportionally impacted by landscape-scale forest clearance and habitat fragmentation, which are prevalent in the study region. Our method also highlights the usefulness of rapid sampling of foraging bats for assaying genetic structure, particularly where roosting sites are not always known.

Wing damage is not used to confirm WNS: In response to elevated prevalence of wing damage in bats near WNS-affected hibernacula in 2008, we proposed a wing damage index (WDI) to objectively monitor wing conditions of bats in the field (Reichard and Kunz, 2009). Francl et al. suggested WDI is used ‘‘...as a proxy for measuring population-level infection’’ with WNS (Francl et al., 2011), but wing damage is only one field sign associated with WNS and does not confirm WNSpositive bats. Instead, WNS is diagnosed by histopathologic confirmation of cutaneous infections attributed to a recently introduced fungus, Geomyces destructans (Meteyer et al., 2009; USGS, 2011). Thus, we argue that suggesting use of WDI as a proxy for infection is inappropriate and misleading.

Natal dispersal is a strategy employed by individuals to avoid reproducing with close relatives. In most bats, incestuous matings are prevented by the departure of one sex; thus, all-offspring philopatry has rarely been recorded in these mammals. Here we present evidence of all-offspring philopatry to natal groups in a Neotropical insectivorous bat. Our results are derived from behavioural observations of a long-term, captureemarkerecapture study conducted on three populations of Spix’s disc-winged bat, Thyroptera tricolor. Results indicate that dispersal patterns of this species are characterized by low emigration rates (0.40e0.73 individuals per year), long residence times (492e1238 days) and high offspring retention of both sexes within natal groups. The retention of offspring resulted in groups being primarily composed of one or two matrilines. Our study is the first to demonstrate such high amounts of male and female natal philopatry coupled with limited spatial movements in bats. These dispersal patterns may be explained by morphological and ecological constraints associated with costly flight dynamics and significant variation in the distribution and quality of habitat, in addition to social benefits accrued by remaining affiliated with close kin. UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biología

The lipid matrix of the stratum corneum (SC), the outer layer of the epidermis of mammals and birds, constitutes the barrier to diffusion of water vapor through the skin. The lipids of the SC are structured in the intercellular spaces of the mammalian epidermis in ordered layers, called lamellae, which have been postulated to prevent water loss. Lipids in the mammalian SC are mainly cholesterol, free fatty acids and ceramides, the latter forming the structural support for the lamellae. However, knowledge on how the lipid composition of the SC alters cutaneous water loss (CWL) in mammals is rudimentary, and is largely derived from studies on laboratory animals and humans. We measured CWL of individuals of two species of syntopic bats, Tadarida brasiliensis and Myotis velifer. In the first study of its kind on wild mammals, we correlated CWL with the lipid composition of the SC, measured using thin layer chromatography and high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. Surface-specific CWL was 20.6% higher in M. velifer than in T. brasiliensis, although differences were not significant. Compared with individuals of M. velifer, individuals of T. brasiliensis had more classes, and a higher proportion, of polar ceramides in the SC, a feature associated with lower CWL. Individuals of T. brasiliensis also had a class of non-polar ceramides that presumably spans the lamellae and gives more cohesiveness to the lipid matrix of the SC. We conclude that qualitative and quantitative modifications of the lipid composition of the SC contribute to regulate CWL of these two species of bats.

Old World leaf-nosed bats (Hipposideridae) are among the most widespread and ecologically diverse groups of insectivorous bats in the Old World tropics. However, phylogenetic relationships in Hipposideridae are poorly resolved at both the generic and species levels, and deep genetic divergence within several Southeast Asian species suggests that current taxonomy underestimates hipposiderid diversity in this region. We used mitochondrial and nuclear sequence data to conduct the first extensive molecular phylogenetic analysis of Southeast Asian hipposiderid bats. Inclusion of multiple samples per taxon allowed testing for evidence of evolutionarily distinct lineages within taxa currently defined as single species. In contrast to earlier phylogenies based on morphometrics, molecular data support monophyly of Hipposideros, but are ambiguous regarding the monophyly of Hipposideridae. With a few exceptions, molecular data also support currently recognized species groups classified by qualitative morphological characters. Widespread paraphyly and polyphyly within many currently recognized species of Hipposideros indicates that evolutionary diversity in the genus is underrepresented by current nomenclature. Comparison of available morphological and echolocation data suggest that both geographic isolation and ecological selection have contributed to the diversification of Southeast Asian hipposiderid bats.

The Brazilian free-tailed bat (Tadarida brasiliensis) exhibits a highly vascularized, hairless thermal window (or “radiator”) on the proximal ventral surfaces of extended wings and body. We identified this character using thermal infrared imaging and investigated the vasculature using barium sulfate enhanced microcomputed tomography (micro-CT). Micro-CT images revealed unique arrangements of arteries and veins in the region of the radiator positioned perpendicular to the axis of the body. Coupling micro-CT imaging with analysis of surface temperature profiles, we concluded that radiators aid in thermoregulation during flight in variable environments. This study represents the first application of contrast enhanced micro-CT to visualize vasculature of bats and thus exhibits a promising technique for further investigations of cardiovascular function and anatomy in bats. Anat Rec, 2012. © 2012 Wiley Periodicals, Inc.

Aeroecology is a discipline that embraces and integrates the domains of atmospheric science, ecology, earth science, geography, computer science, computational biology, and engineering. The unifying concept that underlies this emerging discipline is its focus on the planetary boundary layer, or aerosphere, and the myriad of organisms that, in large part, depend upon this environment for their existence. The aerosphere influences both daily and seasonal movements of organisms, and its effects have both short- and long-term consequences for species that use this environment. The biotic interactions and physical conditions in the aerosphere represent important selection pressures that influence traits such as size and shape of organisms, which in turn facilitate both passive and active displacements. The aerosphere also influences the evolution of behavioral, sensory, metabolic, and respiratory functions of organisms in a myriad of ways. In contrast to organisms that depend strictly on terrestrial or aquatic existence, those that routinely use the aerosphere are almost immediately influenced by changing atmospheric conditions (e.g., winds, air density, precipitation, air temperature), sunlight, polarized light, moon light, and geomagnetic and gravitational forces. The aerosphere has direct and indirect effects on organisms, which often are more strongly influenced than those that spend significant amounts of time on land or in water. Future advances in aeroecology will be made when research conducted by biologists is more fully integrated across temporal and spatial scales in concert with advances made by atmospheric scientists and mathematical modelers. Ultimately, understanding how organisms such as arthropods, birds, and bats aloft are influenced by a dynamic aerosphere will be of importance for assessing, and maintaining ecosystem health, human health, and biodiversity.