The flight of pipistrelle bats Pipistrellus pipistrellus during pregnancy and lactation

A group of 20 pipistrelle bats were taken into captivity and allowed free flight and association within a flight room where they gave birth to and successfully reared 17 young. The flight of the females was recorded during pregnancy, early lactation and post-lactation by using stroboscopic stereophotogrammetry (153 flights reconstructed in total). During the investigation body mass was altering owing to reproductive condition, and changes in mass were recorded daily for all (adult and juvenile) bats during the entire study period, which lasted from two weeks before the last birth until release, when the oldest baby was 43 days old. All bats were individually marked, and detailed morphological measurements were made. Pregnant and post-lactating bats were heavier than lactating bats, which showed the lowest wingbeat frequencies. The flight speeds of pregnant, lactating and post-lactating bats showed no significant differences, and this may be because the pregnant bats appeared to have a wider scope for selecting flight speed than the other two reproductive groups, or than animals studied previously. The group of bats as a whole decreased flight speed (scaling as M^-043) and increased wingbeat frequency (scaling as M^0.58) as their mass increased. Wingbeat amplitude showed no relation to body mass, wing area or span, flight speed or frequency. A flight performance model applied to the experimental results and optimum flight conditions is used to predict cost of transport and mechanical power for steady flight, and equilibrium wingbeat amplitude which is compared with observations.     

Trapped in the darkness of the night: thermal and energetic constraints of daylight flight in bats

Bats are one of the most successful mammalian groups, even though their foraging activities are restricted to the hours of twilight and night-time. Some studies suggested that bats became nocturnal because of overheating when flying in daylight. This is because--in contrast to feathered wings of birds--dark and naked wing membranes of bats efficiently absorb short-wave solar radiation. We hypothesized that bats face elevated flight costs during daylight flights, since we expected them to alter wing-beat kinematics to reduce heat load by solar radiation. To test this assumption, we measured metabolic rate and body temperature during short flights in the tropical short-tailed fruit bat Carollia perspicillata at night and during the day. Core body temperature of flying bats differed by no more than 2°C between night and daytime flights, whereas mass-specific CO(2) production rates were higher by 15 per cent during daytime. We conclude that increased flight costs only render diurnal bat flights profitable when the relative energy gain during daytime is high and risk of predation is low. Ancestral bats possibly have evolved dark-skinned wing membranes to reduce nocturnal predation, but a low degree of reflectance of wing membranes made them also prone to overheating and elevated energy costs during daylight flights. In consequence, bats may have become trapped in the darkness of the night once dark-skinned wing membranes had evolved.     

Wing kinematics of avian flight across speeds

To test whether wing shape affects the kinematics of wing motion during bird flight, we recorded high-speed video (250 Hz) of four species flying in a variable-speed wind tunnel. The birds flew at intervals of 2 m s⁻¹, ranging from 1 m s⁻¹ up to their respective maximum flight speed, which varied from 14 to 17 m s⁻¹ depending on the species. Kinematic data obtained from two synchronized, high-speed video cameras were analyzed using 3D reconstruction. Three species with relatively pointed, high-aspect ratio wings changed wingbeat styles according to flight speed (budgerigar, Melopsittacus undulatus ; cockatiel, Nymphicus hollandicus ; ringed turtle dove, Streptopelia risoria ). These species used a wing-tip reversal upstroke, characterized by supination of the distal wing at mid-upstroke, at equivalent airspeeds ≤7 to 9 m s⁻¹. In faster flight, they used a swept-wing upstroke, without distal wing supination. At mid-upstroke at any speed, wingspan in these species was greater than wrist span. In contrast, at all steady flight speeds, the black-billed magpie Pica hudsonia with relatively broad, low-aspect ratio wings, used a flexed-wing, feathered upstroke in which wrist spans were equal to or greater than wingspans. Our results demonstrate that wing kinematics vary gradually as a function of flight speed, and that the patterns of variation are strongly influenced by external wing shape.     

Perch-hunting in insectivorous Rhinolophus bats is related to the high energy costs of manoeuvring in flight

Foraging behaviour of bats is supposedly largely influenced by the high costs of flapping flight. Yet our understanding of flight energetics focuses mostly on continuous horizontal forward flight at intermediate speeds. Many bats, however, perform manoeuvring flights at suboptimal speeds when foraging. For example, members of the genus Rhinolophus hunt insects during short sallying flights from a perch. Such flights include many descents and ascents below minimum power speed and are therefore considered energetically more expensive than flying at intermediate speed. To test this idea, we quantified the energy costs of short manoeuvring flights (<2 min) using the Na-bicarbonate technique in two Rhinolophus species that differ in body mass but have similar wing shapes. First, we hypothesized that, similar to birds, energy costs of short flights should be higher than predicted by an equation derived for bats at intermediate speeds. Second, we predicted that R. mehelyi encounters higher flight costs than R. euryale, because of its higher wing loading. Although wing loading of R. mehelyi was only 20% larger than that of R. euryale, its flight costs (2.61 ± 0.75 W; mean ± 1 SD) exceeded that of R. euryale (1.71 ± 0.37 W) by 50%. Measured flight costs were higher than predicted for R. mehelyi, but not for R. euryale. We conclude that R. mehelyi face elevated energy costs during short manoeuvring flights due to high wing loading and thus may optimize foraging efficiency by energy-conserving perch-hunting.     

Post-prandial urine loss and its relation to ecology in brown long-eared (Plecotus auritus) and Daubenton's (Myotis daubentoni) bats (Chiroptera: Vespertilionidae)

Urine loss, over the first 12 hours after feeding, was positively and linearly dependent on food consumption in water-denied, brown long-eared bats ( Plecotus auritus ) and also in water-denied and water-provided, Daubenton's bats ( Myotis daubentoni ). The slope of the relationship (food-dependent urine loss) (363 μ 1 g dry mass food⁻¹, S.D.=±70, n=19 ) was not significantly different between the two species but predicted urine loss at zero food consumption (food-independent urine loss) was significantly lower in P. auritus (0.048 μl.min⁻¹, S.D. =±0.015, n= 12) than in M. duubentoni (0.217 μl min⁻¹ S.D. =±0.040, n = 7 ). The same results were apparent if the data for M. daubentoni were restricted to water-deprived animals only. Of total urine loss, 46% occurred in the first hour after feeding in M. daubentoni compared with only 20% in P. auritus . We suggest that the differences between the two species in the pattern of postprandial urine loss reflect their relative association with open water when foraging and roosting in the wild. In the course of the water-denied experiment, M. duubentoni lost 15% of pre-fed body mass and showed signs of severe dehydration, while P. auritus only lost 6% and did not. However, urine loss only accounted for 8–10% of body mass loss. A water budget model for wild P. auritus in the summer was developed and suggested that if bats did not drink, approximately 19% of water loss would be attributable to faecal water loss, 18–20% to urine loss, and 59–62% of intake would be available to support evaporation and reproductive losses.     

Rain increases the energy cost of bat flight

Similar to insects, birds and pterosaurs, bats have evolved powered flight. But in contrast to other flying taxa, only bats are furry. Here, we asked whether flight is impaired when bat pelage and wing membranes get wet. We studied the metabolism of short flights in Carollia sowelli, a bat that is exposed to heavy and frequent rainfall in neotropical rainforests. We expected bats to encounter higher thermoregulatory costs, or to suffer from lowered aerodynamic properties when pelage and wing membranes catch moisture. Therefore, we predicted that wet bats face higher flight costs than dry ones. We quantified the flight metabolism in three treatments: dry bats, wet bats and no rain, wet bats and rain. Dry bats showed metabolic rates predicted by allometry. However, flight metabolism increased twofold when bats were wet, or when they were additionally exposed to rain. We conclude that bats may not avoid rain only because of sensory constraints imposed by raindrops on echolocation, but also because of energetic constraints.     

How swift are swifts Apus apus?

Swifts Apus apus are renowned for their fast flight manner which has fascinated people in all times. However, previous studies of swifts in flight during migration and roosting flights have shown that the birds operate over a narrow range of flight speeds compared with most other birds studied. In this study we have focused on the special flight behavior often called 'screaming parties'. During these flights the birds appear to reach very high speeds and therefore we used a stereo high speed camera setup to measure the flight speeds of the birds during this behavior with high accuracy. The birds were found to fly at much higher speeds during 'screaming parties' than during migration or roosting, on average twice as fast, 20.9  ms⁻¹ (±5.1  ms⁻¹) in horizontal speed. The highest record was 31.1  ms⁻¹ which is the highest measured yet for a swift in self powered flight. Furthermore, the birds were performing steep climbing flights, on average 4.0  ms⁻¹ (±2.8  ms⁻¹) in vertical velocity. A clear trade-off between horizontal speed and vertical speed was found, suggesting that the birds are operating at their maximum.     

Ontogenetic change in the relationship between metabolic rate and body mass in a sea bream Pagrus major (Temminck & Schlegel)

Ontogenetic changes in the relationship between resting rate of oxygen consumption and wet body mass were examined at 20° C with the sea bream Pagrus major ranging from 0.00020 g (weight just after hatching) to 270 g (weight at 530 days old). There was a triphasic relationship between oxygen consumption of an individual fish M (μl min⁻¹) and body mass W (g). During a very early stage (weight 0.00020–0.00025 g), corresponding to the pre-larval stage and with the transitional period to the post-larval stage, there was no substantial change in body mass. The mass–specific metabolic rate M/W (μl g⁻¹ min ¹) showed no clear relationship to body mass as expressed by the equation M/ W =4.86 + 1.47 D , where D is age in days. During the post-larval stage (weight 0.00031–0.005 g), M/W remained almost constant independent of body mass following the expression M = 12.5 W0 .949. During the juvenile and later stages (weight 0.005–270 g), M/ W decreased with increasing body mass following the expression M = 6.3 W ^0.821 which is significantly different from the expression for the post-larval stage ( P < 0.001). Ontogenetic changes in the metabolism-body mass relationship are discussed from the viewpoint of relative growth of organs with different metabolic activities.     

中华山蝠翼的形态学发育模式

为了揭示中华山蝠翼的形态学发育模式,2012-2014年, 通过对50只中华山蝠幼蝠进行人工饲养实验观察及测量,辅以野外标志重捕,研究中华山蝠翼的生长。结果显示：中华山蝠翼外形各量度值先呈直线增长,后增长速度逐渐减慢。各指标增长速度减慢的日龄各不相同,臂膜长在25日龄生长速度逐渐减慢;翼展、翼面积、臂膜面积在30日龄增长速度减慢;掌膜长及掌膜面积则在40日龄增长速度减慢。臂膜长发育最快,试飞前 (28日龄) 已达到成体臂膜长的80.9%。翼载在幼蝠生后14日龄内呈直线增长,然后开始直线下降,到试飞期 (35日龄左右),翼载值降到最低,是成体翼载的80.0%,此后呈极缓慢增长趋势。中华山蝠翼的生长主要集中在试飞前。试飞后,则通过减缓体重的增长速度甚至减轻体重,保持翼面积尤其是掌膜面积的增长速度,从而降低翼载,以便更快地适应飞行生活。Logistic, Von Bertalanffy和Gompertz 3种非线性曲线中,Gompertz模型对翼展拟合度最佳,Gompertz和Von Bertalanffy模型对翼面积拟合度优于Logistic曲线。     

Kinematic plasticity during flight in fruit bats: individual variability in response to loading

All bats experience daily and seasonal fluctuation in body mass. An increase in mass requires changes in flight kinematics to produce the extra lift necessary to compensate for increased weight. How bats modify their kinematics to increase lift, however, is not well understood. In this study, we investigated the effect of a 20% increase in mass on flight kinematics for Cynopterus brachyotis, the lesser dog-faced fruit bat. We reconstructed the 3D wing kinematics and how they changed with the additional mass. Bats showed a marked change in wing kinematics in response to loading, but changes varied among individuals. Each bat adjusted a different combination of kinematic parameters to increase lift, indicating that aerodynamic force generation can be modulated in multiple ways. Two main kinematic strategies were distinguished: bats either changed the motion of the wings by primarily increasing wingbeat frequency, or changed the configuration of the wings by increasing wing area and camber. The complex, individual-dependent response to increased loading in our bats points to an underappreciated aspect of locomotor control, in which the inherent complexity of the biomechanical system allows for kinematic plasticity. The kinematic plasticity and functional redundancy observed in bat flight can have evolutionary consequences, such as an increase potential for morphological and kinematic diversification due to weakened locomotor trade-offs.     

Postnatal development in Andersen's leaf-nosed bat Hipposideros pomona: flight, wing shape, and wing bone lengths

Postnatal changes in flight development, wing shape and wing bone lengths of 56 marked neonate Hipposideros pomona were investigated under natural conditions in southwest China. Flight experiments showed that pups began to flutter with a short horizontal displacement at 10 days and first took flight at 19 days, with most achieving sustained flight at 1 month old. Analysis of covariance on wingspan, wing area, and the other seven wing characteristics between ‘pre-flight’ and ‘post-volancy’ periods supports the hypothesis that growth had one ‘pre-flight’ trajectory and a different ‘post-volancy’ trajectory in bats. Wingspan, handwing length and area, armwing length and area, and total wing area increased linearly until the age of first flight, after which the growth rates decreased (all P < 0.001). Wing loading declined linearly until day 19 before ultimately decreasing to adult levels (P < 0.001). Additionally, the relationship of different pairwise combinations of bony components composing span-wise length and chord-wise length was evaluated to test the hypothesis that compensatory growth of wing bones in H. pomona occurred in both ‘pre-flight’ and ‘post-volancy’ periods. The frequency of short-long and long-short pairs was significantly greater than that of short-short, long-long pairs in most pairs of bone elements in adults. The results indicate that a bone ‘shorter than expected’ would be compensated by a bone or bones ‘longer than expected’, suggesting compensatory growth in H. pomona. The pairwise comparisons conducted in adults were also performed in young bats during ‘pre-flight’ and ‘post-volancy’ periods, demonstrating that compensatory growth occurred throughout postnatal ontogeny.     

Postnatal development in the big-footed bat,Myotis macrodactylus: wing morphology,echolocation calls,and flight

We studied the postnatal development of wing morphology and echolocation calls during flight in a free-ranging population of the big-footed bat, Myotis macrodactylus, using the mark-recapture methodology. Young bats were reluctant to move until 7 days of age and started fluttering at a mean age of 10 days. The wingspan and wing area of pups followed a linear pattern of growth until 22 days of age, by which time the young bats exhibited flapping flight, with mean growth rates of 0.62 mm/day and 3.15 mm²/day, for wingspan and area, respectively, after which growth rates decreased. Pups achieved sustained flight at 40 days of age. Of the three nonlinear growth models (logistic, Gompertz, and von Bertalanffy), the logistic equation provided the best fit to the empirical curves for wingspan and wing area. Neonates emitted long echolocation calls with multiple harmonics. The duration of calls decreased significantly between flutter (19 days) and flight (22 days) stages. The peak and start frequency of calls increased significantly over the 3-week period of development, but the terminal frequency did not change significantly over the development period.     

Determinants of wing-feather moult speed in songbirds

Wing morphology is known to strongly affect flight performance by affecting lift and drag during flight. Performance may consequently deteriorate during feather moult due to the creation of feather gaps in the wing. Since wing gap size may directly affect the extent of reduced flight capacity, rapid moult involving the creation of large feather gaps is expected to substantially impair flight compared with the small gaps induced by a slower moult. To examine the factors affecting wing-feather moult speed, we studied adults of nineteen resident or very short-distance migrant passerine species during their post-breeding moult using a model-selection framework following a phylogenetically controlled analysis. We examined the speed of wing-feather moult in relation to each species’ flight distance index that was estimated based on local foraging movements rather than on longer flights (e.g., migration), assessed by the Delphi technique of expert evaluation. Moult speed was also examined with respect to six morphometric variables: body mass, wing loading, the feather comprising the tip of the wing, aspect ratio, wing span, and wing area. Our results suggest that flight distance index is the most important factor determining the speed of wing-feather moult in songbirds. Species that regularly fly a shorter distance were found to moult quickly, and those that take relatively longer flights moult slowly. These results suggest that the aerodynamic cost of wing area reduction due to feather moult shapes the evolution of annual routine processes by dictating a slower moult speed (resulting in small wing gaps) for species that regularly fly long distances and consequently may be affected more substantially by large wing gaps compared with short distance flyers.     

Intraspecific scaling of flight power in the bat Glossophaga soricina (Phyllostomidae)

Aerodynamic theory predicts that power output during flight should vary with body mass by an exponent of 1.56 when wing morphology remains constant (within an individual), and by an exponent of 1.19 when wing morphology changes with body mass (within a species or between species). I tested these predictions by estimating the power input during horizontal flight in three pregnant and two subadult Glossophaga soricina using a multivariate regression model. This analysis yielded power input during resting and flight as well as the energetic equivalent of change in body mass. A comparison of the estimated flight power for pregnant G. soricina, with published data on flight power of non-pregnant adults, revealed that energy turnover in flight is highest for pregnant G. soricina. Flight power of a 13-g pregnant G. soricina was even higher than that of a 16-g non-pregnant Glossophaga longirostris. A least-squares regression analysis yielded the following equations for the intraspecific scaling of flight power with body mass: power input during horizontal flight (P _f )=24099 body mass (bm; kg)^2.15 (r ²=0.97) for the intra-individual allometry (pregnancy) and P _f =113 bm(kg)^0.95 (r ²=0.99) for the inter-individual allometry (ontogeny). Both mass exponents are not significantly different from the predicted values for the scaling relationship of flight power within an individual (1.56) and within a species (1.19). This is the first measurement of power input during flight for subadult and pregnant bats. Accepted: 11 May 2000     

Development of wing morphology in the Indian pygmy batPipistrellus mimus

The growth and development of the wing parameters of the Indian pygmy batPipistrellus mimus was studied under natural conditions. Newborn young were marked with nontoxic coloured paint and were later marked with split rings. The wingspan and wing area showed linear growth until the age of five weeks, after which the rate of growth decreased. The observations on flight showed that at the age of 19 days the young were able to flutter their wings, at the age of 22 days they flew for a short distance and at the age of 29 days they exhibited sustained flight. The development of wing loading and aspect ratio are also presented. The decrease in wing loading as the bat grows is discussed as an advantage to sustain flight. The aspect ratio showed a high degree of scatter at early stages of life which decreased at the later period of growth. In general the development of wing morphology ofP. mimus is similar to that of other vespertilionid bats.     

Wind as a factor influencing flower-visiting by Hadena bicruris (Noctuidae) and Deilephila elpenor (Sphingidae)

Abstract. 1. In a low-speed wind tunnel, male as well as female moths of Hadena bicruris responded to floral odours with positive anemotaxis. Hitherto, such orientation has only been demonstrated for male moths in response to pheromones.
2. H.bicruris had a maximum flight speed of 4.5–5.4ms^-1 and stopped its flower visiting at a wind speed of 2.5–2.8 m s^-1.
3. Deilephila elpenor had a similar maximum flight speed (4.5— 5.1 ms^-1), but it continued visiting flowers up to wind speeds of 3.0–5.0 ms^-1.
4. Apart from mechanical resistance during flight and flower visits, wind might have adverse effects on the energy budget and on evaporative water loss.     

Patterns of dispersion, density and dispersal in alpine populations of the land snail Arianta arbustorum (L.) (Helicidae)

In studying populations of the land snail Arianta arbustorum in the eastern Swiss Alps, juvenile and subadult snails were found to be aggregated all the time, while adults were mostly aggregated during summer, the main mating period. The mean size of aggregations was 25 cm × 25 cm but varied, according to the area a grass tuft covered, while the intensity of aggregations was influenced by weather conditions. Densities of 10–15 subadults and 3–6 adults m⁻² were found along ditches and streams, 2–25 subadults and 2–20 adults on uncultivated alpine meadows, while on scree-covered alpine grassland the density of subadults and adults was 0.3–0.5 m⁻². The extent of daily movements varied with microclimatic factors and season and depended on the structure of the habitat. The dispersal rate was greatest in June (2.6 m month⁻¹) and least in August (1.1 m month⁻¹). In summer and autumn, active uphill movements were noted, a process which compensates for the loss of altitude by passive downhill displacements. Roiling down on snowfields and being carried away by avalanches and streams were observed.     

Flight style in bats as predicted from wing morphometry: the effects of specimen preservation

An as yet unconsidered potential error in studies that predict flight style from morphological measurements of bats is the effect of the specimen type employed. On the basis of the finding that morphological measurements taken from fluid-preserved bat specimens may not yield values equivalent to those taken from the live animal, we compared the values of several variables (lifting surface area, wingspan, mass, aspect ratio, wing loading and minimum power speed) for live and fluid-preserved little brown bats ( Myotis lucifugus ) with the accepted standards for this species given by Norberg & Rayner (1987). Significant differences were detected for lifting surface area, wingspan, mass, aspect ratio and wing loading values taken from live bats and their respective values reported by Norberg & Rayner. Differences between preserved bats and Norberg & Rayner's numbers were limited to lifting surface area and wingspan (extended wing positions only), aspect ratio (all wing positions), and mass (both 70% ethanol- and 45% isopropyl alcohol-preserved specimens). Thus, Norberg & Rayner's values correspond most closely to values obtained from preserved museum specimens, a fact reflecting the source of their data in this instance. This and other limitations involved in attempting to predict the flight style of bats from a few morphological characters are discussed.     

Effect of Iysozyme concentration, heating at 90°C, and then incubation at chilled temperatures on growth from spores of non-proteolytic Clostridium botulinum

The heat treatment necessary to inactivate spores of non-proteolytic Clostridium botulinum in refrigerated, processed foods may be influenced by the occurrence of lysozyme in these foods. Spores of six strains of non-proteolytic Cl. botulinum were inoculated into tubes of an anaerobic meat medium, to give 10⁶ spores per tube. Hen egg white lysozyme (0–50 μg ml^-1) was added, and the tubes were given a heat treatment equivalent to 19·8 min at 90°C, cooled, and incubated at 8°, 12°, 16° and 25°C for up to 93 d. In the absence of added lysozyme, neither growth nor toxin formation were observed. A 6–D inactivation was therefore achieved. In tubes to which lysozyme (5–50 μg ml^-1) had been added prior to heating, growth and toxin formation were observed. With lysozyme added at 50 μg ml^-1, growth was first observed after 68 d at 8°C, 31 d at 12°C, 24 d at 16°C, and 9 d at 25°C. Thus, in these circumstances, a heat treatment equivalent to 19·8 min at 90°C was not sufficient, on its own, to give a 6–D inactivation. A combination of the heat treatment, maintenance at less than 12°C, and a shelf-life not more than 4 weeks reduced the risk of growth of non-proteolytic Cl. botulinum by a factor of 10⁶.