Detection of prey in a cluttered environment by the northern bat Eptesicus nilssonii.

We studied the acoustic behaviour of bats (Eptesicus nilssonii) hunting for large (wing span 5 cm) non-hearing hepialid moths (Hepialus humuli). Groups of silvery-white male H. humuli perform a short (30 min) hovering display flight over grassland at dusk. They typically hover at, or below, the tops of grass panicles and are therefore situated in a highly acoustically cluttered habitat. Occasionally, they move to a new position by making short (1-5 s) flights at higher levels. E. nilssonii is not a clutter specialist, and yet we found that they attacked H. humuli within the 'clutter overlap zone'. The bats did not change their signal design in any marked manner for this specific task. Measurements of echoes from a moth 10 cm above or below the grass tops showed that information for detecting the moths was available to the bats. Nevertheless, the bats did not attack moths in stationary hovering display flight, only when they moved above the grass panicles. The duration of the up/down flights (movements) were almost always longer than an entire capture sequence by the bats. Apparently, the bats rely on the movement of the moth in space, monitored over successive echoes, to discriminate moth echoes from overlapping clutter echoes.

Persistence of bat defence reactions in high Arctic moths (Lepidoptera).

We investigated the bat defence reactions of three species of moths (Gynaephora groenlandica, Gynaephora rossi (Lymantriidae) and Psychophora sabini (Geometridae)) in the Canadian Arctic archipelago. Since these moths inhabit the Arctic tundra and, therefore, are most probably spatially isolated from bats, their hearing and associated defensive reactions are probably useless and would therefore be expected to disappear with ongoing adaptation to Arctic conditions. When exposed to bat-like ultrasound (26 kHz and 110 dB sound pressure level root mean square at 1 m) flying male Gynaephora spp. always reacted defensively by rapidly reversing their flight course. They could hear the sound and reacted at least 15-25 m away. Psychophora sabini walking on a surface froze at distances of at least 5-7 m from the sound source. However, two out of three individuals of this species (all males) did not respond in any way to the sound while in flight. Hence, we found evidence of degeneration of bat defence reactions, i.e. adaptation to the bat-free environment, in P. sabini but not in Gynaephora spp. Some Arctic moths (Gynaephora spp.) still possess defensive reactions against bats, possibly because the selection pressure for the loss of the trait is such that it declines only very slowly (perhaps by genetic drift; and there may not have been enough time for the trait to disappear. One possible reason may be that Arctic moths have long generation times.

Bat defence in lekking ghost swifts (Hepialus humuli), a moth without ultrasonic hearing.

The Hepialidae represents an early branch of the Lepidoptera, whose members lack the ultrasonic hearing and other obvious predator defence systems present in other extant moths. I observed lekking male ghost swifts, Hepialus humuli, being exploited by northern bats, Eptesicus nilssonii, over a hayfield in southern Sweden. Because the moth's display flight was restricted to a brief (30 min) period at dusk, they avoided most predators temporally but were exposed to early emerging aerial-hawking bats. Against these, they apparently employed 'acoustic crypsis', achieved by flying close (< 0.5 m) to the vegetation, thereby hiding from the bats among clutter (echoes returning from the background). Nevertheless, the predation risk for the displaying moth males was very high (20% per night), mainly because they sometimes left the safety of the vegetation. The lack of 'advanced' predator defence mechanisms in H. humuli requires alternative defence strategies, which, however, restrict the behavioural repertoire and still carry a high predation risk.

Mercury vapour lamps interfere with the bat defence of tympanate moths (Operophtera spp.; Geometridae)

Bats often forage near streetlamps, where they catch moths in particular. At least two hypotheses may explain the apparent increase in the availability of moths to bats feeding around streetlamps: (1) the moths become concentrated near the light and therefore more profitable to exploit; and (2) the light interferes with the moths' evasive flight behaviour. We tested the second of these hypotheses by exposing flying male winter moths, Operophtera spp., to bursts of ultrasound (26 kHz, 110 dB sound pressure level) from an electronic source. The light from a 125 W mercury vapour lamp had a quantitative effect on the moths' evasive flight response at close range (within ca 4 m), inhibiting it totally in nearly half (43%, N=125) of the cases. By contrast, moths flying in the surrounding woodland and without interference from the lamp always responded to the sound. Streetlamps of the mercury vapour type (white lamps) thus interfere with the defensive behaviour of moths and presumably increase their vulnerability to echolocating bats. This may have implications for the conservation of both moths and bats. Copyright 1998 The Association for the Study of Animal Behaviour.

Hearing and bat defence in geometrid winter moths.

Audiograms and behavioural responses to ultrasound reveal that male geometrid winter moths (Agriopis and Erannis spp.; Ennominae, and Alsophila aescularia; Oenochrominae), which have large wings and a slow flight, have good, broadly tuned ultrasonic hearing with best frequencies at 25-40 kHz, coinciding with the frequencies used by most sympatric aerial-hawking bats. Ultrasonic pulses (27 kHz 110 dB at 1 m) delivered at distances of 1-12 m evoked consistent reactions of free flying, male A. marginaria in the lab as well as in the field; those at < 5 m resulted in the moth spiralling or diving towards the ground, those at 5-12 m resulted in one or several changes in the flight path, but did not end on the ground. The differential reaction probably reflects whether the moth is likely to have been detected by the bat or not. The micropterous (and flightless), and hence cryptic, females have strongly reduced tympanic organs and are virtually deaf. Sexual dimorphism in hearing and behavioural reactions to ultrasound reflect differential natural selection on males and females by bats. Natural selection on the hearing of the males thus seems to occur although they fly in late autumn and early spring, when bat activity is much reduced.

Primary focal segmental glomerular sclerosis in adults: prognostic value of histologic variants.

Primary focal segmental glomerular sclerosis (FSGS) is a clinicopathologic syndrome in which variable amounts of proteinuria are associated with the renal biopsy finding of segmental glomerular scarring in some, but not all, of the glomeruli. Additional histologic features have been described in FSGS, including the position of the scar relative to the vascular and tubular pole of the glomerulus, foam cells, hyalinosis, mesangial deposits of immunoglobulin M, diffuse mesangial hypercellularity, glomerular visceral epithelial cell hyperplasia and hypertrophy, and the extent of associated interstitial fibrosis and tubular atrophy. We performed a retrospective study on 81 patients with biopsy-proven, primary FSGS to determine whether any of the histologic features of FSGS correlated with renal function at the time of biopsy and the incidence of end-stage renal disease at follow-up. Sixty patients were nephrotic and 21 had nonnephrotic proteinuria. Only the degree of interstitial fibrosis correlated with the initial serum creatinine (r = 0.536) and none of the histologic features predicted the presence of nephrotic-range proteinuria at the time of biopsy. Segmental scars involved 21% +/- 14% of the glomeruli per biopsy specimen, but their position within the glomerulus was uniform in only 13% of the cases. Diffuse mesangial hypercellularity was present in 17% of the biopsy specimens, and glomerular epithelial cell lesions were present in 57% of the biopsy specimens. Multivariate analysis showed that only the degree of interstitial fibrosis predicted end-stage renal disease in all 81 patients and in the 60 patients with nephrotic-range proteinuria. The current data do not support different therapeutic approaches in primary FSGS based on histologic subtypes.

Low-frequency echolocation enables the bat Tadarida teniotis to feed on tympanate insects.

The European free-tailed bat, Tadarida teniotis, forages in uncluttered airspace by using intense narrow-band echolocation calls with low frequency (11-12 kHz), and feeds on relatively large flying insects, mainly (90% by volume) of the tympanate orders Lepidoptera and Neuroptera. The use of low-frequency echolocation calls without strong harmonics appears to be a specialization for long-range detection of large, tympanate insects, which are less well represented in the diet of most other aerial-hawking bats. The results provide evidence in support of the allotonic frequency hypothesis, i.e. that use of echolocation calls with frequencies above or below the best hearing of tympanate insects is an adaptation to increase the availability of these insects.