Species biomass and size composition of fish faunas along a productivity gradient were studied in south Swedish lakes. Generally, with increasing productivity (measured as chlorophyll content), Salmoniformes were replaced by percids, which in turn were replaced by cyprinids, as suggested in previous studies. However, percids showed two peaks in biomass, one in medium productive lakes due to perch and one in highly productive lakes due to zander. Benthic piscivores were present in all lakes, whereas pelagic piscivores were absent in the least productive lakes. The proportion of piscivores in the total fish biomass showed a peak in medium productive lakes, largely reflecting the importance of piscivorous perch. The median size of the dominant cyprinids (roach) in the systems studied increased as the importance of piscivores increased, which was interpreted as a size refuge response to increased predation pressure. The same pattern was present for the dominant planktivore, vendace, in low productive systems. Although a predictable pattern ofchange in the fish fauna was found along the productivity gradient, other environmental factors such as structural complexity may be the ultimate cause of the observed succession pattern.
Staying aloft when hovering and flying slowly is demanding. According to quasi-steady-state aerodynamic theory, slow-flying vertebrates should not be able to generate enough lift to remain aloft. Therefore, unsteady aerodynamic mechanisms to enhance lift production have been proposed. Using digital particle image velocimetry, we showed that a small nectar-feeding bat is able to increase lift by as much as 40% using attached leading-edge vortices (LEVs) during slow forward flight, resulting in a maximum lift coefficient of 4.8. The airflow passing over the LEV reattaches behind the LEV smoothly to the wing, despite the exceptionally large local angles of attack and wing camber. Our results show that the use of unsteady aerodynamic mechanisms in flapping flight is not limited to insects but is also used by larger and heavier animals.
The flapping flight of animals generates an aerodynamic footprint as a time-varying vortex wake in which the rate of momentum change represents the aerodynamic force. We showed that the wakes of a small bat species differ from those of birds in some important respects. In our bats, each wing generated its own vortex loop. Also, at moderate and high flight speeds, the circulation on the outer (hand) wing and the arm wing differed in sign during the upstroke, resulting in negative lift on the hand wing and positive lift on the arm wing. Our interpretations of the unsteady aerodynamic performance and function of membranous-winged, flapping flight should change modeling strategies for the study of equivalent natural and engineered flying devices.
Many of the morphological features of animals are considered to be adaptations to the habitat that the animals utilize. The habitats utilized by birds vary, perhaps more than for any other group of vertebrates. Here, we study possible adaptations in the morphology of the skeletal elements of the hind limbs to the habitat of birds. Measurements of the lengths of the femur, tibiotarsus and tarsometatarsus of 323 bird species from 74 families are used together with body mass data, taken from the literature. The species are separated into six habitat groups on the basis of literature data on leg use. A discriminant analysis of the groups based on leg morphology shows that swimming birds, wading birds and ground living species are more easily identified than other birds. Furthermore, functional predictions are made for each group based on ecological and mechanical considerations. The groups were tested for deviation from the norm for all birds for three indices of size‐ and leg‐length‐independent measures of the bones and for a size‐independent‐index of leg length. Several of the groups deviate significantly from the norm for one or more of the indices used, suggesting habitat‐related adaptations in the leg morphology of birds. The results indicate that stability is an important factor affecting the leg morphology of primarily long‐legged birds. The femur seems to be more important than previously thought because several of the groups have high femur indices, suggesting a positive selection pressure on this bone. On a general basis, the results suggest that the effect of leg length should be taken into consideration when discussing adaptations of mass‐independent lengths of the long bones of the legs of birds. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 79, 461–484.
SUMMARY To obtain a full understanding of the aerodynamics of animal flight, the movement of the wings, the kinematics, needs to be connected to the wake left behind the animal. Here the detailed 3D wingbeat kinematics of bats, Glossophaga soricina, flying in a wind tunnel over a range of flight speeds (1–7 m s−1) was determined from high-speed video. The results were compared with the wake geometry and quantitative wake measurements obtained simultaneously to the kinematics. The wingbeat kinematics varied gradually with flight speed and reflected the changes observed in the wake of the bats. In particular, several of the kinematic parameters reflected the differences in the function of the upstroke at low and high flight speeds. At lower flight speeds the bats use a pitch-up rotation to produce a backward flick which creates thrust and some weight support. At higher speeds this mechanism disappears and the upstroke generates weight support but no thrust. This is reflected by the changes in e.g. angle of attack, span ratio, camber and downstroke ratio. We also determined how different parameters vary throughout a wingbeat over the flight speeds studied. Both the camber and the angle of attack varied over the wingbeat differently at different speeds, suggesting active control of these parameters to adjust to the changing aerodynamic conditions. This study of the kinematics strongly indicates that the flight of bats is governed by an unsteady high-lift mechanism at low flight speeds and points to differences between birds and bats.
Bats are unique among extant actively flying animals in having very flexible wings, controlled by multi-jointed fingers. This gives the potential for fine-tuned active control to optimize aerodynamic performance throughout the wingbeat and thus a more efficient flight. But how bat wing performance scales with size, morphology and ecology is not yet known.Here, we present time-resolved fluid wake data of two species of bats flying freely across a range of flight speeds using stereoscopic digital particle image velocimetry in a wind tunnel. From these data, we construct an average wake for each bat species and speed combination, which is used to estimate the flight forces throughout the wingbeat and resulting flight performance properties such as lift-to-drag ratio (L/D). The results show that the wake dynamics and flight performance of both bat species are similar, as was expected since both species operate at similar Reynolds numbers (Re) and Strouhal numbers (St). However, maximum L/D is achieved at a significant higher flight speed for the larger, highly mobile and migratory bat species than for the smaller non-migratory species. Although the flight performance of these bats may depend on a range of morphological and ecological factors, the differences in optimal flight speeds between the species could at least partly be explained by differences in their movement ecology.
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