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.
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