Diurnal and nocturnal African dung beetles use celestial cues, such as the sun, the moon, and the polarization pattern, to roll dung balls along straight paths across the savanna. Although nocturnal beetles move in the same manner through the same environment as their diurnal relatives, they do so when light conditions are at least 1 million-fold dimmer. Here, we show, for the first time to our knowledge, that the celestial cue preference differs between nocturnal and diurnal beetles in a manner that reflects their contrasting visual ecologies. We also demonstrate how these cue preferences are reflected in the activity of compass neurons in the brain. At night, polarized skylight is the dominant orientation cue for nocturnal beetles. However, if we coerce them to roll during the day, they instead use a celestial body (the sun) as their primary orientation cue. Diurnal beetles, however, persist in using a celestial body for their compass, day or night. Compass neurons in the central complex of diurnal beetles are tuned only to the sun, whereas the same neurons in the nocturnal species switch exclusively to polarized light at lunar light intensities. Thus, these neurons encode the preferences for particular celestial cues and alter their weighting according to ambient light conditions. This flexible encoding of celestial cue preferences relative to the prevailing visual scenery provides a simple, yet effective, mechanism for enabling visual orientation at any light intensity.he blue sky is a rich source of visual cues that are used by many animals during orientation or navigation (1, 2). Besides the sun, celestial phenomena, such as the skylight intensity gradient or the more complex polarization pattern, can serve as references for spatial orientation (3-5). Polarized skylight is generated by scattered sunlight in the atmosphere, and to a terrestrial observer, the resulting alignment of the electric field vectors extends across the entire sky, forming concentric circles around the position of the sun (Fig. 1A). A similar distribution of brightness and polarization pattern is also created around the moon (6). Although this nocturnal pattern is 1 million-fold dimmer than the daylight pattern (6), some animals, such as South African ball-rolling dung beetles, can use this lunar polarization pattern for orientation (7). To avoid competition for food at the dung pile, these beetles detach a piece of dung, shape it into a ball, and roll it away along a straightline path. For this type of straight-line orientation, nocturnal beetles seem to rely exclusively on celestial cues (8), such as the moon or polarized light.As with all nocturnal animals, night-active beetles have to overcome a major challenge: They need to maintain high orientation precision even under extremely dim light conditions. Indeed, recent experiments have shown that nocturnal dung beetles orient at night with the same precision as their diurnal relatives during the day (9), an ability partly due to the fact that their eyes are considerably more sensiti...

We explore the degeneracy between mass and spin in gravitational waveforms emitted by black-hole binary coalescences. We focus on spin-aligned waveforms and obtain our results using phenomenological models that were tuned to numerical-relativity simulations. A degeneracy is known for low-mass binaries (particularly neutron-star binaries), where gravitational-wave detectors are sensitive to only the inspiral phase, and the waveform can be modelled by post-Newtonian theory. Here, we consider black-hole binaries, where detectors will also be sensitive to the merger and ringdown, and demonstrate that the degeneracy persists across a broad mass range. At low masses, the degeneracy is between mass ratio and total spin, with chirp mass accurately determined. At higher masses, the degeneracy persists but is not so clearly characterised by constant chirp mass as the merger and ringdown become more significant. We consider the importance of this degeneracy both for performing searches (including searches where only non-spinning templates are used) and in parameter extraction from observed systems. We compare observational capabilities between the early (~2015) and final (2018 onwards) versions of the Advanced LIGO detector.Comment: 11 pages, 9 figure

Global expansion of human activities is associated with the introduction of novel stimuli, such as anthropogenic noise, artificial lights, and chemical agents. Progress in documenting the ecological effects of sensory pollutants is weakened by sparse knowledge of the mechanisms underlying these effects. This severely limits our capacity to devise mitigation measures. Here, we integrate knowledge of animal sensory ecology, physiology, and life history to articulate three perceptual mechanismsmasking, distracting, and misleadingthat clearly explain how and why anthropogenic sensory pollutants impact organisms. We then link these three mechanisms to ecological consequences, and discuss their implications for conservation. We argue that this framework can reveal the presence of 'sensory danger zones', hotspots of conservation concern where sensory pollutants overlap in space and time with an organism's activity, and foster development of strategic interventions to mitigate the impact of sensory pollutants. Future research that applies this framework will provide critical insight to preserve the natural sensory world.

In order to protect their food from competitors, ball-rolling dung beetles detach a piece of dung from a pile, shape it into a ball, and roll it away along a straight path [1]. They appear to rely exclusively on celestial compass cues to maintain their bearing [2-8], but the mechanism that enables them to use these cues for orientation remains unknown. Here, we describe the orientation strategy that allows dung beetles to use celestial cues in a dynamic fashion. We tested the underlying orientation mechanism by presenting beetles with a combination of simulated celestial cues (sun, polarized light, and spectral cues). We show that these animals do not rely on an innate prediction of the natural geographical relationship between celestial cues, as other navigating insects seem to [9, 10]. Instead, they appear to form an internal representation of the prevailing celestial scene, a "celestial snapshot," even if that scene represents a physical impossibility for the real sky. We also find that the beetles are able to maintain their bearing with respect to the presented cues only if the cues are visible when the snapshot is taken. This happens during the "dance," a behavior in which the beetle climbs on top of its ball and rotates about its vertical axis [11]. This strategy for reading celestial signals is a simple but efficient mechanism for straight-line orientation.

To escape competition at the dung pile, a ball-rolling dung beetle forms a piece of dung into a ball and rolls it away. To ensure their efficient escape from the dung pile, beetles rely on a 'celestial compass' to move along a straight path. Here, we analyzed the reliability of different skylight cues for this compass and found that dung beetles rely not only on the sun but also on the skylight polarization pattern. Moreover, we show the first evidence of an insect using the celestial light-intensity gradient for orientation. Using a polarizer, we manipulated skylight so that the polarization pattern appeared to turn by 90 deg. The beetles then changed their bearing close to the expected 90 deg. This behavior was abolished if the sun was visible to the beetle, suggesting that polarized light is hierarchically subordinate to the sun. When the sky was depolarized and the sun was invisible, the beetles could still move along straight paths. Therefore, we analyzed the use of the celestial light-intensity gradient for orientation. Artificial rotation of the intensity pattern by 180 deg caused beetles to orient in the opposite direction. This lightintensity cue was also found to be subordinate to the sun and could play a role in disambiguating the polarization signal, especially at low sun elevations.

The quality of visual information that is available to an animal is limited by the size of its eyes. Differences in eye size can be observed even between closely related individuals but we understand little about how this affects visual quality. Insects are good models for exploring the effects of size on visual systems because many species exhibit size polymorphism, which modifies both the size and shape of their eyes. Previous work in this area has been limited, however, due to the challenge of determining the 3D structure of eyes. To address this, we have developed a novel method based on x-ray tomography to measure the 3D structure of insect eyes and calculate their visual capabilities. We investigated visual allometry in the bumblebee Bombus terrestris and found that size affects specific aspects of visual quality including binocular overlap, optical sensitivity across the field of view, and visual resolution in the dorsofrontal visual field. This holistic study on eye allometry reveals that differential scaling between different eye areas provides substantial flexibility for larger bumblebees to have improved visual capabilities. List of abbreviations:az. -Azimuth CC -Crystalline cone el. -Elevation EV -Eye volume FOV -Field of view ITW -Inter-tegula width IO -Inter-ommatidial microCT -micro-computed tomography NV -Normal vector

To control flight, insects rely on the pattern of visual motion generated on the retina as they move through the environment. When light levels fall, vision becomes less reliable and flight control thus becomes more challenging. Here, we investigated the effect of light intensity on flight control by filming the trajectories of free-flying bumblebees (Bombus terrestris, Linnaeus 1758) in an experimental tunnel at different light levels. As light levels fell, flight speed decreased and the flight trajectories became more tortuous but the bees were still remarkably good at centring their flight about the tunnel's midline. To investigate whether this robust flight performance can be explained by visual adaptations in the bumblebee retina, we also examined the response speed of the green-sensitive photoreceptors at the same light intensities. We found that the response speed of the photoreceptors significantly decreased as light levels fell. This indicates that bumblebees have both behavioural (reduction in flight speed) and retinal (reduction in response speed of the photoreceptors) adaptations to allow them to fly in dim light. However, the more tortuous flight paths recorded in dim light suggest that these adaptations do not support flight with the same precision during the twilight hours of the day.

Visually guided flight control in the rainforest is arguably one of the most complex insect behaviors: illumination varies dramatically depending on location [1], and the densely cluttered environment blocks out most of the sky [2]. What visual information do insects sample for flight control in this habitat? To begin answering this question, we determined the visual fields of the ocelli-thought to play a role in attitude stabilization of some flying insects [3-5]-of an orchid bee, Euglossa imperialis. High-resolution 3D models of the ocellar system from X-ray microtomography were used for optical ray tracing simulations. Surprisingly, these showed that each ocellus possesses two distinct visual fields-a focused monocular visual field suitable for detecting features elevated above the horizon and therefore assisting with flight stabilization [3-5] and, unlike other ocelli investigated to date [4, 6, 7], a large trinocular fronto-dorsal visual field shared by all ocelli. Histological analyses show that photoreceptors have similar orientations within each ocellus and are likely to be sensitive to polarized light, as in some other hymenopterans [7, 8]. We also found that the average receptor orientation is offset between the ocelli, each having different axes of polarization sensitivity relative to the head. Unlike the eyes of any other insect described to date, this ocellar system meets the requirements of a true polarization analyzer [9, 10]. The ocelli of E. imperialis could provide sensitive compass information for navigation in the rainforest and, additionally, provide cues for visual discrimination or flight control.