Miklós Blahó

Polarotactic tabanids find striped patterns with brightness and/or polarization modulation least attractive: an advantage of zebra stripes.

SUMMARY The 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 zebra’s 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.

An unexpected advantage of whiteness in horses: the most horsefly-proof horse has a depolarizing white coat

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 hosts 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 tabanid–horse interaction, our results can probably be extrapolated to other host animals of polarotactic tabanids, as the reflection–polarization characteristics of the hosts body surface are physically the same, and thus not species-dependent.

Reducing the Maladaptive Attractiveness of Solar Panels to Polarotactic Insects

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.

New kind of polarotaxis governed by degree of polarization: attraction of tabanid flies to differently polarizing host animals and water surfaces

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.

No evidence for behavioral responses to circularly polarized light in four scarab beetle species with circularly polarizing exocuticle

The strongest known circular polarization of biotic origin is the left-circularly polarized (LCP) light reflected from the metallic shiny exocuticle of certain beetles of the family Scarabaeidae. This phenomenon has been discovered by Michelson in 1911. Although since 1955 it has been known that the human eye perceives a visual illusion when stimulated by circularly polarized (CP) light, it was discovered only recently that a stomatopod shrimp is able to perceive circular polarization. It is pertinent to suppose that scarab beetles reflecting LCP light in an optical environment (vegetation) being deficient in CP signals may also perceive circular polarization and use it to find each other (mate/conspecifics) as until now it has been believed. We tested this hypothesis in six choice experiments with several hundred individuals of four scarab species: Anomala dubia, Anomala vitis (Coleoptera, Scarabaeidae, Rutelinae), and Cetonia aurata, Potosia cuprea (Coleoptera, Scarabaeidae, Cetoniinae), all possessing left-circularly polarizing exocuticle. From the results of our experiments we conclude that the studied four scarab species are not attracted to CP light when feeding or looking for mate or conspecifics. We demonstrated that the light reflected by host plants of the investigated scarabs is circularly unpolarized. Our results finally solve a puzzle raised over one hundred years ago, when Michaelson discovered that scarab beetles reflect circularly polarized light.

Spottier targets are less attractive to tabanid flies: on the tabanid-repellency of spotty fur patterns.

During blood-sucking, female members of the family Tabanidae transmit pathogens of serious diseases and annoy their host animals so strongly that they cannot graze, thus the health of the hosts is drastically reduced. Consequently, a tabanid-resistant coat with appropriate brightness, colour and pattern is advantageous for the host. Spotty coats are widespread among mammals, especially in cattle (Bos primigenius). In field experiments we studied the influence of the size and number of spots on the attractiveness of test surfaces to tabanids that are attracted to linearly polarized light. We measured the reflection-polarization characteristics of living cattle, spotty cattle coats and the used test surfaces. We show here that the smaller and the more numerous the spots, the less attractive the target (host) is to tabanids. We demonstrate that the attractiveness of spotty patterns to tabanids is also reduced if the target exhibits spottiness only in the angle of polarization pattern, while being homogeneous grey with a constant high degree of polarization. Tabanid flies respond strongly to linearly polarized light, and we show that bright and dark parts of cattle coats reflect light with different degrees and angles of polarization that in combination with dark spots on a bright coat surface disrupt the attractiveness to tabanids. This could be one of the possible evolutionary benefits that explains why spotty coat patterns are so widespread in mammals, especially in ungulates, many species of which are tabanid hosts.

Stripes disrupt odour attractiveness to biting horseflies: Battle between ammonia, CO2, and colour pattern for dominance in the sensory systems of host-seeking tabanids

As with mosquitoes, female tabanid flies search for mammalian hosts by visual and olfactory cues, because they require a blood meal before being able to produce and lay eggs. Polarotactic tabanid flies find striped or spotted patterns with intensity and/or polarisation modulation visually less attractive than homogeneous white, brown or black targets. Thus, this reduced optical attractiveness to tabanids can be one of the functions of striped or spotty coat patterns in ungulates. Ungulates emit CO2 via their breath, while ammonia originates from their decaying urine. As host-seeking female tabanids are strongly attracted to CO2 and ammonia, the question arises whether the poor visual attractiveness of stripes and spots to tabanids is or is not overcome by olfactory attractiveness. To answer this question we performed two field experiments in which the attractiveness to tabanid flies of homogeneous white, black and black-and-white striped three-dimensional targets (spheres and cylinders) and horse models provided with CO2 and ammonia was studied. Since tabanids are positively polarotactic, i.e. attracted to strongly and linearly polarised light, we measured the reflection-polarisation patterns of the test surfaces and demonstrated that these patterns were practically the same as those of real horses and zebras. We show here that striped targets are significantly less attractive to host-seeking female tabanids than homogeneous white or black targets, even when they emit tabanid-luring CO2 and ammonia. Although CO2 and ammonia increased the number of attracted tabanids, these chemicals did not overcome the weak visual attractiveness of stripes to host-seeking female tabanids. This result demonstrates the visual protection of striped coat patterns against attacks from blood-sucking dipterans, such as horseflies, known to transmit lethal diseases to ungulates.

How can horseflies be captured by solar panels? A new concept of tabanid traps using light polarization and electricity produced by photovoltaics

Horseflies (Diptera: Tabanidae) can cause severe problems for humans and livestock because of the continuous annoyance performed and the diseases vectored by the haematophagous females. Therefore, effective horsefly traps are in large demand, especially for stock-breeders. To catch horseflies, several kinds of traps have been developed, many of them attracting these insects visually with the aid of a black ball. The recently discovered positive polarotaxis (attraction to horizontally polarized light) in several horsefly species can be used to design traps that capture female and male horseflies. The aim of this work is to present the concept of such a trap based on two novel principles: (1) the visual target of the trap is a horizontal solar panel (photovoltaics) attracting polarotactic horseflies by means of the highly and horizontally polarized light reflected from the photovoltaic surface. (2) The horseflies trying to touch or land on the photovoltaic trap surface are perished by the mechanical hit of a wire rotated quickly with an electromotor supplied by the photovoltaics-produced electricity. Thus, the photovoltaics is bifunctional: its horizontally polarized reflected light signal attracts water-seeking, polarotactic horseflies, and it produces the electricity necessary to rotate the wire. We describe here the concept and design of this new horsefly trap, the effectiveness of which was demonstrated in field experiments. The advantages and disadvantages of the trap are discussed. Using imaging polarimetry, we measured the reflection-polarization characteristics of the photovoltaic trap surface demonstrating the optical reason for the polarotactic attractiveness to horseflies.

Phototaxis and polarotaxis hand in hand: night dispersal flight of aquatic insects distracted synergistically by light intensity and reflection polarization

Based on an earlier observation in the field, we hypothesized that light intensity and horizontally polarized reflected light may strongly influence the flight behaviour of night-active aquatic insects. We assumed that phototaxis and polarotaxis together have a more harmful effect on the dispersal flight of these insects than they would have separately. We tested this hypothesis in a multiple-choice field experiment using horizontal test surfaces laid on the ground. We offered simultaneously the following visual stimuli for aerial aquatic insects: (1) lamplit matte black canvas inducing phototaxis alone, (2) unlit shiny black plastic sheet eliciting polarotaxis alone, (3) lamplit shiny black plastic sheet inducing simultaneously phototaxis and polarotaxis, and (4) unlit matte black canvas as a visually unattractive control. The unlit matte black canvas trapped only a negligible number (13) of water insects. The sum (16,432) of the total numbers of water beetles and bugs captured on the lamplit matte black canvas (7,922) and the unlit shiny black plastic sheet (8,510) was much smaller than the total catch (29,682) caught on the lamplit shiny black plastic sheet. This provides experimental evidence for the synergistic interaction of phototaxis (elicited by the unpolarized direct lamplight) and polarotaxis (induced by the strongly and horizontally polarized plastic-reflected light) in the investigated aquatic insects. Thus, horizontally polarizing artificial lamplit surfaces can function as an effective ecological trap due to this synergism of optical cues, especially in the urban environment.

A horizontally polarizing liquid trap enhances the tabanid-capturing efficiency of the classic canopy trap

Host-seeking female tabanid flies, that need mammalian blood for the development of their eggs, can be captured by the classic canopy trap with an elevated shiny black sphere as a luring visual target. The design of more efficient tabanid traps is important for stock-breeders to control tabanids, since these blood-sucking insects can cause severe problems for livestock, especially for horse- and cattle-keepers: reduced meat/milk production in cattle farms, horses cannot be ridden, decreased quality of hides due to biting scars. We show here that male and female tabanids can be caught by a novel, weather-proof liquid-filled black tray laid on the ground, because the strongly and horizontally polarized light reflected from the black liquid surface attracts water-seeking polarotactic tabanids. We performed field experiments to reveal the ideal elevation of the liquid trap and to compare the tabanid-capturing efficiency of three different traps: (1) the classic canopy trap, (2) the new polarization liquid trap, and (3) the combination of the two traps. In field tests, we showed that the combined trap captures 2.4-8.2 times more tabanids than the canopy trap alone. The reason for the larger efficiency of the combined trap is that it captures simultaneously the host-seeking female and the water-seeking male and female tabanids. We suggest supplementing the traditional canopy trap with the new liquid trap in order to enhance the tabanid-capturing efficiency.