The alteration of natural cycles of light and dark by artificial light sources has deleterious impacts on animals and ecosystems. Many animals can also exploit a unique characteristic of light – its direction of polarization –as a source of information. We introduce the term “polarized light pollution” (PLP) to focus attention on the ecological consequences of light that has been polarized through interaction with human‐made objects. Unnatural polarized light sources can trigger maladaptive behaviors in polarization‐sensitive taxa and alter ecological interactions. PLP is an increasingly common byproduct of human technology, and mitigating its effects through selective use of building materials is a realistic solution. Our understanding of how most species use polarization vision is limited, but the capacity of PLP to drastically increase mortality and reproductive failure in animal populations suggests that PLP should become a focus for conservation biologists and resource managers alike.
SUMMARYThe characteristic striped appearance of zebras has provoked much speculation about its function and why the pattern has evolved, but experimental evidence is scarce. Here, we demonstrate that a zebra-striped horse model attracts far fewer horseflies (tabanids) than either homogeneous black, brown, grey or white equivalents. Such biting flies are prevalent across Africa and have considerable fitness impact on potential mammalian hosts. Besides brightness, one of the likely mechanisms underlying this protection is the polarization of reflected light from the host animal. We show that the attractiveness of striped patterns to tabanids is also reduced if only polarization modulations (parallel stripes with alternating orthogonal directions of polarization) occur in horizontal or vertical homogeneous grey surfaces. Tabanids have been shown to respond strongly to linearly polarized light, and we demonstrate here that the light and dark stripes of a zebraʼs coat reflect very different polarizations of light in a way that disrupts the attractiveness to tabanids. We show that the attractiveness to tabanids decreases with decreasing stripe width, and that stripes below a certain size are effective in not attracting tabanids. Further, we demonstrate that the stripe widths of zebra coats fall in a range where the striped pattern is most disruptive to tabanids. The striped coat patterns of several other large mammals may also function in reducing exposure to tabanids by similar mechanisms of differential brightness and polarization of reflected light. This work provides an experimentally supported explanation for the underlying mechanism leading to the selective advantage of a black-and-white striped coat pattern. Supplementary material available online at
White horses frequently suffer from malign skin cancer and visual deficiencies owing to their high sensitivity to the ultraviolet solar radiation. Furthermore, in the wild, white horses suffer a larger predation risk than dark individuals because they can more easily be detected. In spite of their greater vulnerability, white horses have been highly appreciated for centuries owing to their natural rarity. Here, we show that blood-sucking tabanid flies, known to transmit disease agents to mammals, are less attracted to white than dark horses. We also demonstrate that tabanids use reflected polarized light from the coat as a signal to find a host. The attraction of tabanids to mainly black and brown fur coats is explained by positive polarotaxis. As the host's colour determines its attractiveness to tabanids, this parameter has a strong influence on the parasite load of the host. Although we have studied only the tabanidhorse interaction, our results can probably be extrapolated to other host animals of polarotactic tabanids, as the reflection-polarization characteristics of the host's body surface are physically the same, and thus not species-dependent.
Ecological photopollution created by artificial night lighting can alter animal behavior and lead to population declines and biodiversity loss. Polarized light pollution is a second type of photopollution that triggers water-seeking insects to ovisposit on smooth and dark man-made objects, because they simulate the polarization signatures of natural water bodies. We document a case study of the interaction of these two forms of photopollution by conducting observations and experiments near a lamp-lit bridge over the river Danube that attracts mass swarms of the mayfly Ephoron virgo away from the river to oviposit on the asphalt road of the bridge. Millions of mayflies swarmed near bridge-lights for two weeks. We found these swarms to be composed of 99% adult females performing their upstream compensatory flight and were attracted upward toward unpolarized bridge-lamp light, and away from the horizontally polarized light trail of the river. Imaging polarimetry confirmed that the asphalt surface of the bridge was strongly and horizontally polarized, providing a supernormal ovipositional cue to Ephoron virgo, while other parts of the bridge were poor polarizers of lamplight. Collectively, we confirm that Ephoron virgo is independently attracted to both unpolarized and polarized light sources, that both types of photopollution are being produced at the bridge, and that spatial patterns of swarming and oviposition are consistent with evolved behaviors being triggered maladaptively by these two types of light pollution. We suggest solutions to bridge and lighting design that should prevent or mitigate the impacts of such scenarios in the future. The detrimental impacts of such scenarios may extend beyond Ephoron virgo.
Human-made objects (e.g., buildings with glass surfaces) can reflect horizontally polarized light so strongly that they appear to aquatic insects to be bodies of water. Insects that lay eggs in water are especially attracted to such structures because these insects use horizontal polarization of light off bodies of water to find egg-laying sites. Thus, these sources of polarized light can become ecological traps associated with reproductive failure and mortality in organisms that are attracted to them and by extension with rapid population declines or collapse. Solar panels are a new source of polarized light pollution. Using imaging polarimetry, we measured the reflection-polarization characteristics of different solar panels and in multiple-choice experiments in the field we tested their attractiveness to mayflies, caddis flies, dolichopodids, and tabanids. At the Brewster angle, solar panels polarized reflected light almost completely (degree of polarization d ≈ 100%) and substantially exceeded typical polarization values for water (d ≈ 30-70%). Mayflies (Ephemeroptera), stoneflies (Trichoptera), dolichopodid dipterans, and tabanid flies (Tabanidae) were the most attracted to solar panels and exhibited oviposition behavior above solar panels more often than above surfaces with lower degrees of polarization (including water), but in general they avoided solar cells with nonpolarizing white borders and white grates. The highly and horizontally polarizing surfaces that had nonpolarizing, white cell borders were 10- to 26-fold less attractive to insects than the same panels without white partitions. Although solar panels can act as ecological traps, fragmenting their solar-active area does lessen their attractiveness to polarotactic insects. The design of solar panels and collectors and their placement relative to aquatic habitats will likely affect populations of aquatic insects that use polarized light as a behavioral cue.
Adult tabanid flies (horseflies and deerflies) are terrestrial and lay their eggs onto marsh plants near bodies of fresh water because the larvae develop in water or mud. To know how tabanids locate their host animals, terrestrial rendezvous sites and egg-laying places would be very useful for control measures against them, because the hematophagous females are primary/secondary vectors of some severe animal/human diseases/parasites. Thus, in choice experiments performed in the field we studied the behavior of tabanids governed by linearly polarized light. We present here evidence for positive polarotaxis, i.e., attraction to horizontally polarized light stimulating the ventral eye region, in both males and females of 27 tabanid species. The novelty of our findings is that positive polarotaxis has been described earlier only in connection with the water detection of some aquatic insects ovipositing directly into water. A further particularity of our discovery is that in the order Diptera and among blood-sucking insects the studied tabanids are the first known species possessing ventral polarization vision and definite polarization-sensitive behavior with known functions. The polarotaxis in tabanid flies makes it possible to develop new optically luring traps being more efficient than the existing ones based on the attraction of tabanids by the intensity and/or color of reflected light.
Aquatic insects find their habitat from a remote distance by means of horizontal polarization of light reflected from the water surface. This kind of positive polarotaxis is governed by the horizontal direction of polarization (E-vector). Tabanid flies also detect water by this kind of polarotaxis. The host choice of blood-sucking female tabanids is partly governed by the linear polarization of light reflected from the host's coat. Since the coat-reflected light is not always horizontally polarized, host finding by female tabanids may be different from the established horizontal E-vector polarotaxis. To reveal the optical cue of the former polarotaxis, we performed choice experiments in the field with tabanid flies using aerial and ground-based visual targets with different degrees and directions of polarization. We observed a new kind of polarotaxis being governed by the degree of polarization rather than the E-vector direction of reflected light. We show here that female and male tabanids use polarotaxis governed by the horizontal E-vector to find water, while polarotaxis based on the degree of polarization serves host finding by female tabanids. As a practical by-product of our studies, we explain the enigmatic attractiveness of shiny black spheres used in canopy traps to catch tabanids.
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