Blood-sucking female tabanid flies cause serious problems for animals and humans. For the control of tabanids, the knowledge about their seasonality and daily activity is of great importance. Earlier, only traditional traps capturing exclusively female tabanids have been used to survey tabanid activity. The data of such temporal trapping do not reflect correctly the activity of male and female tabanid flies. Our major aim was to monitor the trapping numbers of male and female tabanids during a 3-month summer survey in Hungary. We used (i) conventional canopy traps with liquid traps on the ground beneath the canopy and (ii) L-shaped sticky traps with vertical and horizontal components. Our other goal was to compare the efficiencies of the two components of each trap type used. We observed two greater peaks of the trapping number of tabanids. These peaks started with increased catches of female tabanids captured by the canopy traps and the vertical sticky traps and ended with a dominance of male and female tabanids caught by the liquid traps and the horizontal sticky traps. The swarming periods were interrupted by rainy/cool days, when the number of tabanids decreased drastically. Among the 17 species, six dominated and composed 89.4 % of the captured tabanids: Haematopota pluvialis, Tabanus tergestinus, Tabanus bromius, Tabanus maculicornis, Tabanus bovinus and Atylotus loewianus. The number of water-seeking male and female tabanids rose up to 12-13 h and then decreased but had a secondary peak at about 17 h. The stochastic weather change and the communities of different species resulted in large standard deviations of the averaged number of tabanids in the course of a day. The horizontally polarizing (liquid and horizontal sticky) traps captured both male and female specimens and were about three times more efficient than the canopy and vertical sticky traps that caught only females. The results of the horizontal sticky traps corresponded to those of the liquid traps, while the catches of the vertical sticky traps corresponded to those of the canopy traps. The catches of the used trap types reflected well the species and water/host-seeking composition of tabanids.
It is a widely discussed hypothesis that Viking seafarers might have been able to locate the position of the occluded sun by means of dichroic or birefringent crystals, the mysterious sunstones, with which they could analyze skylight polarization. Although the atmospheric optical prerequisites and certain aspects of the efficiency of this sky-polarimetric Viking navigation have been investigated, the accuracy of the main steps of this method has not been quantitatively examined. To fill in this gap, we present here the results of a planetarium experiment in which we measured the azimuth and elevation errors of localization of the invisible sun. In the planetarium sun localization was performed in two selected celestial points on the basis of the alignments of two small sections of two celestial great circles passing through the sun. In the second step of sky-polarimetric Viking navigation the navigator needed to determine the intersection of two such celestial circles. We found that the position of the sun (solar elevation ?(S), solar azimuth ?(S)) was estimated with an average error of +0.6°????+8.8° and -3.9°????+2.0°. We also calculated the compass direction error when the estimated sun position is used for orienting with a Viking sun-compass. The northern direction (?(North)) was determined with an error of -3.34°???(North)?+6.29°. The inaccuracy of the second step of this navigation method was high (??(North)=-16.3°) when the solar elevation was 5°??(S)?25°, and the two selected celestial points were far from the sun (at angular distances 95°??(1), ?(2)?115°) and each other (125°???145°). Considering only this second step, the sky-polarimetric navigation could be more accurate in the mid-summer period (June and July), when in the daytime the sun is high above the horizon for long periods. In the spring (and autumn) equinoctial period, alternative methods (using a twilight board, for example) might be more appropriate. Since Viking navigators surely also committed further errors in the first and third steps, the orientation errors presented here underestimate the net error of the whole sky-polarimetric navigation.
Vikings routinely crossed the North Atlantic without a magnetic compass and left their mark on lands as far away as Greenland, Newfoundland and Baffin Island. Based on an eleventh-century dial fragment artefact, found at Uunartoq in Greenland, it is widely accepted that they sailed along chosen latitudes using primitive Sun compasses. Such instruments were tested on sea and proved to be efficient hand-held navigation tools, but the dimensions and incisions of the Uunartoq find are far from optimal in this role. On the basis of the sagas mentioning sunstones, incompatible hypotheses were formed for Viking solar navigation procedures and primitive skylight polarimetry with dichroic or birefringent crystals. We describe here a previously unconceived method of navigation based on the Uunartoq artefact functioning as a 'twilight board', which is a combination of a horizon board and a Sun compass optimized for use when the Sun is close to the horizon. We deduced an appropriate solar navigation procedure using a twilight board, a shadow-stick and birefringent crystals, which bring together earlier suggested methods in harmony and provide a true skylight compass function. This could have allowed Vikings to navigate around the clock, to use the artefact dial as a Sun compass during long parts of the day and to use skylight polarization patterns in the twilight period. In field tests, we found that true north could be appointed with such a medieval skylight compass with an error of about ±4° when the artificially occluded Sun had elevation angles between +10° and -8° relative to the horizon. Our interpretation allows us to assign exact dates to the gnomonic lines on the artefact and outlines the schedule of the merchant ships that sustained the Viking colony in Greenland a millennium ago.
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.
The horizontally polarizing surface parts of shiny black cars (the reflection-polarization characteristics of which are similar to those of water surfaces) attract water-leaving polarotactic insects. Thus, shiny black cars are typical sources of polarized light pollution endangering water-leaving insects. A new fashion fad is to make car-bodies matt black or grey. Since rough (matt) surfaces depolarize the reflected light, one of the ways of reducing polarized light pollution is to make matt the concerned surface. Consequently, matt black/grey cars may not induce polarized light pollution, which would be an advantageous feature for environmental protection. To test this idea, we performed field experiments with horizontal shiny and matt black car-body surfaces laid on the ground. Using imaging polarimetry, in multiple-choice field experiments we investigated the attractiveness of these test surfaces to various water-leaving polarotactic insects and obtained the following results: (i) The attractiveness of black car-bodies to polarotactic insects depends in complex manner on the surface roughness (shiny, matt) and species (mayflies, dolichopodids, tabanids). (ii) Non-expectedly, the matt dark grey car finish is much more attractive to mayflies (being endangered and protected in many countries) than matt black finish. (iii) The polarized light pollution of shiny black cars usually cannot be reduced with the use of matt painting. On the basis of these, our two novel findings are that (a) matt car-paints are highly polarization reflecting, and (b) these matt paints are not suitable to repel polarotactic insects. Hence, the recent technology used to make matt the car-bodies cannot eliminate or even can enhance the attractiveness of black/grey cars to water-leaving insects. Thus, changing shiny black car painting to matt one is a disadvantageous fashion fad concerning the reduction of polarized light pollution of black vehicles.
It is widely accepted that Vikings used sun-compasses to derive true directions from the cast shadow of a gnomon. It has been hypothesized that when a cast shadow was not formed, Viking navigators relied on crude skylight polarimetry with the aid of dichroic or birefringent crystals, called "sunstones." We demonstrate here that a simple tool, that we call "shadow-stick," could have allowed orientation by a sun-compass with satisfying accuracy when shadows were not formed, but the sun position could have reliably been estimated. In field tests, we performed orientation trials with a set composed of a sun-compass, two calcite sunstones, and a shadow-stick. We show here that such a set could have been an effective orientation tool for Vikings only when clear, blue patches of the sky were visible.
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.
Trapping flies with sticky paper sheets is an ancient method. The classic flypaper has four typical characteristics: (i) its sticky paper is bright (chamois, light yellow or white), (ii) it is strip-shaped, (iii) it hangs vertically, and (iv) it is positioned high (several metres) above ground level. Such flypapers, however, do not trap horseflies (tabanids). There is a great need to kill horseflies with efficient traps because they are vectors of dangerous diseases, and due to their continuous annoyance livestock cannot graze, horses cannot be ridden, and meat and milk production from cattle is drastically reduced. Based on earlier findings on the positive polarotaxis (attraction to linearly polarised light) in tabanid flies and modifying the concept of the old flypaper, we constructed a new horsefly trap called "horseflypaper". In four field experiments we showed that the ideal horseflypaper (i) is shiny black, (ii) has an appropriately large (75×75 cm(2)) surface area, (iii) has sticky black vertical and horizontal surfaces in an L-shaped arrangement, and (iv) its horizontal surface should be at ground level for maximum effectiveness. Using imaging polarimetry, we measured the reflection-polarisation characteristics of this new polarisation tabanid trap. The ideal optical and geometrical characteristics of this trap revealed in field experiments are also explained. The horizontal part of the trap captures water-seeking male and female tabanids, while the vertical part catches host-seeking female tabanids.
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.
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.
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.
The experts of animal locomotion well know the characteristics of quadruped walking since the pioneering work of Eadweard Muybridge in the 1880s. Most of the quadrupeds advance their legs in the same lateral sequence when walking, and only the timing of their supporting feet differ more or less. How did this scientific knowledge influence the correctness of quadruped walking depictions in the fine arts? Did the proportion of erroneous quadruped walking illustrations relative to their total number (i.e. error rate) decrease after Muybridge? How correctly have cavemen (upper palaeolithic Homo sapiens) illustrated the walking of their quadruped prey in prehistoric times? The aim of this work is to answer these questions. We have analyzed 1000 prehistoric and modern artistic quadruped walking depictions and determined whether they are correct or not in respect of the limb attitudes presented, assuming that the other aspects of depictions used to determine the animals gait are illustrated correctly. The error rate of modern pre-Muybridgean quadruped walking illustrations was 83.5%, much more than the error rate of 73.3% of mere chance. It decreased to 57.9% after 1887, that is in the post-Muybridgean period. Most surprisingly, the prehistoric quadruped walking depictions had the lowest error rate of 46.2%. All these differences were statistically significant. Thus, cavemen were more keenly aware of the slower motion of their prey animals and illustrated quadruped walking more precisely than later artists.
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.
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 hosts 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.
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.
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 zebras 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.
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