Wavelength-specific negatively phototactic responses of the burrowing mayfly larvae Ephoron virgo.

Mayflies are typically negatively phototactic during larval development, while their adults possess positive phototaxis. However, no extensive research was made about the wavelength dependence of phototaxis in any mayfly larvae. We measured the repellency rate of Ephoron virgo larvae to light as a function of wavelength in the 368 - 743 nm spectral range. We established that the magnitude of repellence increased with decreasing wavelength and the maximal responses were elicited by 400 nm violet light. This wavelength dependence of phototaxis is similar to the recently reported spectral sensitivity of positive phototaxis of the twilight-swarming E. virgo adults. Negative phototaxis not only facilitates evading predation, avoidance of the blue-violet spectral range could promote the larvae to withdraw towards the river midline in the case of drop in the water level, when the underwater light becomes enriched with shorter wavelengths due to the decreasing depth of overhead river water.

Spectral sensitivity transition in the compound eyes of a twilight-swarming mayfly and its visual ecological implications.

Aquatic insect species that leave the water after larval development, such as mayflies, have to deal with extremely different visual environments in their different life stages. Measuring the spectral sensitivity of the compound eyes of the virgin mayfly (Ephoron virgo) resulted in differences between the sensitivity of adults and larvae. Larvae were primarily green-, while adults were mostly UV-sensitive. The sensitivity of adults and larvae was the same in the UV, but in the green spectral range, adults were 3.3 times less sensitive than larvae. Transmittance spectrum measurements of larval skins covering the eye showed that the removal of exuvium during emergence cannot explain the spectral sensitivity change of the eyes. Taking numerous sky spectra from the literature, the ratio of UV and green photons in the skylight was shown to be maximal for θ ≈ -13° solar elevation, which is in the θmin = -14.7° and θmax = -7.1° typical range of swarming that was established from webcam images of real swarmings. We suggest that the spectral sensitivity of both the larval and adult eyes are adapted to the optical environment of the corresponding life stages.

Polarization neutral point pairs of the solar corona and the lunar disc observed during the total solar eclipse on 11 August 1999 in Hungary.

During the total solar eclipses on 11 August 1999 in Kecel (Hungary) and on 29 March 2006 in Side (Turkey), two Hungarian groups performed full-sky imaging polarimetric measurements of the eclipsed sky. They observed the spatiotemporal change of the celestial polarization pattern and detected three polarization neutral points as well as two points with maximal polarization of the sky. Parallel to these studies, the polarization pattern in front of the lunar disc, the solar corona, and the surrounding sky have also been measured. During the total solar eclipse on 21 August 2017 in the USA (Rexburg-Idaho, Madras-Oregon), three American/international groups have measured the polarization characteristics of the full sky and the solar corona. The first group observed the spatiotemporal variation of the celestial polarization pattern, and the second group detected three polarization neutral points of the sky and observed two neutral points of the solar corona. The latter were named as Minnaert and van de Hulst neutral points. The third group observed two neutral points of the lunar disc. We have reanalyzed the earlier polarization patterns of the lunar disc, solar corona, and the surrounding sky measured during the Hungarian total eclipse on 11 August 1999. In these reanalyzed polarization patterns, all four neutral points observed during the eclipse on 21 August 2017 in the USA occur: the Minnaert/van de Hulst neutral point pair above/below the eclipsed Sun, where coronal polarization is canceled by sky polarization, and the northern and southern (unnamed) neutral points of the lunar disc, where the directions of polarization of coronal light and foreground skylight are perpendicular to each other with the same polarized intensity. We name the latter two polarization neutral points after Coulson and Vorobiev.

Bioreplicated coatings for photovoltaic solar panels nearly eliminate light pollution that harms polarotactic insects.

Many insect species rely on the polarization properties of object-reflected light for vital tasks like water or host detection. Unfortunately, typical glass-encapsulated photovoltaic modules, which are expected to cover increasingly large surfaces in the coming years, inadvertently attract various species of water-seeking aquatic insects by the horizontally polarized light they reflect. Such polarized light pollution can be extremely harmful to the entomofauna if polarotactic aquatic insects are trapped by this attractive light signal and perish before reproduction, or if they lay their eggs in unsuitable locations. Textured photovoltaic cover layers are usually engineered to maximize sunlight-harvesting, without taking into consideration their impact on polarized light pollution. The goal of the present study is therefore to experimentally and computationally assess the influence of the cover layer topography on polarized light pollution. By conducting field experiments with polarotactic horseflies (Diptera: Tabanidae) and a mayfly species (Ephemeroptera: Ephemera danica), we demonstrate that bioreplicated cover layers (here obtained by directly copying the surface microtexture of rose petals) were almost unattractive to these species, which is indicative of reduced polarized light pollution. Relative to a planar cover layer, we find that, for the examined aquatic species, the bioreplicated texture can greatly reduce the numbers of landings. This observation is further analyzed and explained by means of imaging polarimetry and ray-tracing simulations. The results pave the way to novel photovoltaic cover layers, the interface of which can be designed to improve sunlight conversion efficiency while minimizing their detrimental influence on the ecology and conservation of polarotactic aquatic insects.

Horsefly reactions to black surfaces: attractiveness to male and female tabanids versus surface tilt angle and temperature.

Tabanid flies (Diptera: Tabanidae) are attracted to shiny black targets, prefer warmer hosts against colder ones and generally attack them in sunshine. Horizontally polarised light reflected from surfaces means water for water-seeking male and female tabanids. A shiny black target above the ground, reflecting light with high degrees and various directions of linear polarisation is recognised as a host animal by female tabanids seeking for blood. Since the body of host animals has differently oriented surface parts, the following question arises: How does the attractiveness of a tilted shiny black surface to male and female tabanids depend on the tilt angle δ? Another question relates to the reaction of horseflies to horizontal black test surfaces with respect to their surface temperature. Solar panels, for example, can induce horizontally polarised light and can reach temperatures above 55 °C. How long times would horseflies stay on such hot solar panels? The answer of these questions is important not only in tabanid control, but also in the reduction of polarised light pollution caused by solar panels. To study these questions, we performed field experiments in Hungary in the summer of 2019 with horseflies and black sticky and dry test surfaces. We found that the total number of trapped (male and female) tabanids is highest if the surface is horizontal (δ = 0°), and it is minimal at δ = 75°. The number of trapped males decreases monotonously to zero with increasing δ, while the female catch has a primary maximum and minimum at δ = 0° and δ = 75°, respectively, and a further secondary peak at δ = 90°. Both sexes are strongly attracted to nearly horizontal (0° ≤ δ ≤ 15°) surfaces, and the vertical surface is also very attractive but only for females. The numbers of touchdowns and landings of tabanids are practically independent of the surface temperature T. The time period of tabanids spent on the shiny black horizontal surface decreases with increasing T so that above 58 °C tabanids spent no longer than 1 s on the surface. The horizontally polarised light reflected from solar panels attracts aquatic insects. This attraction is adverse, if the lured insects lay their eggs onto the black surface and/or cannot escape from the polarised signal and perish due to dehydration. Using polarotactic horseflies as indicator insects in our field experiment, we determined the magnitude of polarised light pollution (being proportional to the visual attractiveness to tabanids) of smooth black oblique surfaces as functions of δ and T.

How does the water springtail optically locate suitable habitats? Spectral sensitivity of phototaxis and polarotaxis in Podura aquatica.

Optical detection of horizontally polarized light is widespread among aquatic insects. This process usually occurs in the UV or blue spectral ranges. Recently, it was demonstrated that at least one collembolan species, the water springtail (Podura aquatica) also possesses positive polarotaxis to horizontally polarized light. These hexapods are positively phototactic, live on the surface of calm waters and usually accumulate close to the riparian vegetation. In laboratory experiments, we measured the wavelength dependence of phototaxis and polarotaxis of P. aquatica in the 346-744 nm and 421-744 nm ranges, respectively. According to our results, the action spectrum of phototaxis is bimodal with two peaks in the blue (λ1=484 nm) and green-yellow (λ2=570 nm) ranges, while polarotaxis operates in the blue spectral range. For the first time, we show that collembolan polarotaxis functions in the same spectral range as the polarotaxis of many aquatic insects. We present our experiments and discuss the possible ecological significance of our findings.

Striped bodypainting protects against horseflies.

Bodypainting is widespread in African, Australian and Papua New Guinean indigenous communities. Many bodypaintings use white or bright yellow/grey/beige stripes on brown skin. Where the majority of people using bodypainting presently live, blood-sucking horseflies are abundant, and they frequently attack the naked brown regions of the human body surface with the risk of transmitting the pathogens of dangerous diseases. Since horseflies are deterred by the black and white stripes of zebras, we hypothesized that white-striped paintings on dark brown human bodies have a similar effect. In a field experiment in Hungary, we tested this hypothesis. We show that the attractiveness to horseflies of a dark brown human body model significantly decreases, if it is painted with the white stripes that are used in bodypaintings. Our brown human model was 10 times more attractive to horseflies than the white-striped brown model, and a beige model, which was used as a control, attracted two times more horseflies than the striped brown model. Thus, white-striped bodypaintings, such as those used by African and Australian people, may serve to deter horseflies, which is an advantageous byproduct of these bodypaintings that could lead to reduced irritation and disease transmission by these blood-sucking insects.

Method to reduce motion artifacts of sequential imaging polarimetry: long enough exposures minimize polarization blurs of wavy water surfaces.

Researchers studying the polarization characteristics of the optical environment prefer to use sequential imaging polarimetry, because it is inexpensive and simple. This technique takes polarization pictures through polarizers in succession. Its main drawback is, however, that during sequential exposure of the polarization pictures, the target must not move, otherwise so-called motion artifacts are caused after evaluation of the polarization pictures. How could these disturbing motion artifacts be minimized? Taking inspiration from photography, our idea was to take the polarization pictures with an exposure that is long enough so that the changes of the moving/changing target can be averaged and, thus, motion artifacts are reduced, at least in a special case when the motion has a stable mean. In the laboratory, we demonstrated the performance of this method when the target was a wavy water surface. We found that the errors of the measured degree and angle of polarization of light reflected from the undulating water surface decreased with increasing exposure time (shutter speed) and converged to very low values. Although various simultaneous polarimeters (taking the polarization pictures at once) are available that do not suffer from motion artifacts, our method is much cheaper and performs very well, at least when the target is a wavy water surface.

Method to improve the survival of night-swarming mayflies near bridges in areas of distracting light pollution.

Numerous negative ecological effects of urban lighting have been identified during the last decades. In spite of the development of lighting technologies, the detrimental effect of this form of light pollution has not declined. Several insect species are affected including the night-swarming mayfly Ephoron virgo: when encountering bridges during their mass swarming, these mayflies often fall victim to artificial lighting. We show a simple method for the conservation of these mayflies exploiting their positive phototaxis. With downstream-facing light-emitting diode beacon lights above two tributaries of the river Danube, we managed to guide egg-laying females to the water and prevent them from perishing outside the river near urban lights. By means of measuring the mayfly outflow from the river as a function of time and the on/off state of the beacons, we showed that the number of mayflies exiting the river's area was practically zero when our beacons were operating. Tributaries could be the sources of mayfly recolonization in case of water quality degradation of large rivers. The protection of mayfly populations in small rivers and safeguarding their aggregation and oviposition sites is therefore important.

Polarization sensitivity in Collembola: an experimental study of polarotaxis in the water-surface-inhabiting springtail Podura aquatica.

The ventral eye of the water-surface-inhabiting springtail Podura aquatica has six ommatidia with horizontal and vertical microvilli and perceives light from the ventral, frontal and frontodorsal regions, whereas the dorsal eye possesses two upward-looking ommatidia with vertical microvilli. The ventral eye may detect water by its polarization sensitivity, even if the insect is resting with its head slightly tipped down on a raised surface. The polarization sensitivity and polarotaxis in springtails (Collembola) have not been investigated. Therefore, we performed behavioural choice experiments to study them in P. aquatica We found that the strength of phototaxis in P. aquatica depends on the polarization characteristics of stimulating light. Horizontally and vertically polarized light were the most and least attractive, respectively, while unpolarized stimulus elicited moderate attraction. We show that horizontally polarized light attracts more springtails than unpolarized, even if the polarized stimulus was 10 times dimmer. Thus, besides phototaxis, P. aquatica also performs polarotaxis with the ability to measure or at least estimate the degree of polarization. Our results indicate that the threshold d* of polarization sensitivity in P. aquatica is between 10.1 and 25.5%.

Mayflies are least attracted to vertical polarization: A polarotactic reaction helping to avoid unsuitable habitats.

Like other aquatic insects, mayflies are positively polarotactic and locate water surfaces by means of the horizontal polarization of water-reflected light. However, may vertically polarized light also have implications for the swarming behaviour of mayflies? To answer this question, we studied in four field experiments the behavioural responses of Ephoron virgo and Caenis robusta mayflies to lamps emitting horizontally and vertically polarized and unpolarized light. In both species, unpolarized light induces positive phototaxis, horizontally polarized light elicits positive photo- and polarotaxis, horizontally polarized light is much more attractive than unpolarized light, and vertically polarized light is the least attractive if the stimulus intensities and spectra are the same. Vertically polarized light was the most attractive for C. robusta if its intensity was about two and five times higher than that of the unpolarized and horizontally polarized stimuli, respectively. We suggest that the mayfly behaviour observed in our experiments may facilitate the stability of swarming above water surfaces. Beside the open water surface reflecting horizontally polarized light, the shadow and mirror image of riparian vegetation at the edge of the water surface reflect weakly and non-horizontally (mainly vertically) polarized light. Due to their positive polarotaxis, flying mayflies remain continuously above the water surface, because they keep away from the unpolarized or non-horizontally polarizing edge regions (water surface and coast line) of water bodies. We also discuss how our findings can explain the regulation of mayfly colonization.

Lamp-lit bridges as dual light-traps for the night-swarming mayfly, Ephoron virgo: interaction of polarized and unpolarized light pollution.

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.

Seasonality and daily activity of male and female tabanid flies monitored in a Hungarian hill-country pasture by new polarization traps and traditional canopy traps.

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.

Unexpected attraction of polarotactic water-leaving insects to matt black car surfaces: mattness of paintwork cannot eliminate the polarized light pollution of black cars.

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.

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.

Orientation with a Viking sun-compass, a shadow-stick, and two calcite sunstones under various weather conditions.

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.

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.

A new tabanid trap applying a modified concept of the old flypaper: linearly polarising sticky black surfaces as an effective tool to catch polarotactic horseflies.

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.

Cavemen were better at depicting quadruped walking than modern artists: erroneous walking illustrations in the fine arts from prehistory to today.

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.

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.