Speedy bearings to slacked steering: Mapping the navigation patterns and motions of Viking voyages.

Viking sailors ruled the North Atlantic Ocean for about three hundred years. Their main sailing route was the 60° 21' 55'' latitude between Norway and Greenland. Although they did not have a magnetic compass, in sunshine they used a sun-compass to determine the geographical north (solar Viking navigation: SVN). It has been hypothesized that when the Sun was invisible, Viking navigators determined the direction of polarization of skylight with sunstones (dichroic/birefringent crystals), and then estimated the geographical north using the sun-compass (sky-polarimetric Viking navigation: SPVN). Many details of the hypothetical SPVN have been thoroughly revealed in psychophysical laboratory and planetarium experiments. Combining these results with measured celestial polarization patterns, the success of SPVN was obtained as functions of sailing, meteorological and navigation parameters (sunstone type, sailing date, navigation periodicity, night sailing, cloudiness conditions). What was so far lacking in this experimental and computational archeological approach is the study of the success of SVN and a combined navigation using solar cues in sunshine (SVN) and sky polarization at invisible Sun (SPVN), the latter being the most realistic method. In this work we determine the success of the sole SVN and the combined SVN-SPVN relative to the mere SPVN for three navigator types (determining the intended sailing direction with large, medium or small frequencies) at spring equinox and summer solstice, with and without night sailing. We found that to maximize the sailing success, navigators had to choose different navigation methods depending on the navigation frequency. Using sky polarization with very frequent navigation, resulted in the highest chance to survive a three-week voyage from Norway to Greenland.

Sensitivity and robustness of sky-polarimetric Viking navigation: Sailing success is most sensitive to night sailing, navigation periodicity and sailing date, but robust against weather conditions.

Although Viking sailors did not have a magnetic compass, they could successfully navigate with a sun-compass under a sunny sky. Under cloudy/foggy conditions, they might have applied the sky-polarimetric Viking navigation (SPVN), the high success of which has been demonstrated with computer simulations using the following input data: sky polarization patterns measured with full-sky imaging polarimetry, and error functions of the navigation steps measured in psychophysical laboratory and planetarium experiments. As a continuation of the earlier studies, in this work we investigate the sensitivity of the success of SPVN to the following relevant sailing, meteorological and navigational parameters: sunstone type, sailing date, navigation periodicity, night sailing, dominance of strongly, medium or weakly cloudy skies, and changeability of cloudiness. Randomly varying these parameters in the simulation of Viking voyages along the latitude 60° 21' 55'' N from Norway to Greenland, we determined those parameters which had strong and weak influences on the success of SPVN. The following intrinsic parameters of the simulation were also randomly changed: sailing speed, visibility distance of Greenland's southeast coastline and start time of diurnal sailing. Our results show that the sailing success is sensitive to the night sailing, navigation periodicity and sailing date, while it is robust against the sunstone type, dominance of strongly, medium or weakly cloudy skies, and changing cloudiness.

A new argument against cooling by convective air eddies formed above sunlit zebra stripes.

There is a long-lasting debate about the possible functions of zebra stripes. According to one hypothesis, periodical convective air eddies form over sunlit zebra stripes which cool the body. However, the formation of such eddies has not been experimentally studied. Using schlieren imaging in the laboratory, we found: downwelling air streams do not form above the white stripes of light-heated smooth or hairy striped surfaces. The influence of stripes on the air stream formation (facilitating upwelling streams and hindering horizontal stream drift) is negligible higher than 1-2 cm above the surface. In calm weather, upwelling air streams might form above sunlit zebra stripes, however they are blown off by the weakest wind, or even by the slowest movement of the zebra. These results forcefully contradict the thermoregulation hypothesis involving air eddies.

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.

Why do biting horseflies prefer warmer hosts? tabanids can escape easier from warmer targets.

Blood-sucking horseflies (tabanids) prefer warmer (sunlit, darker) host animals and generally attack them in sunshine, the reason for which was unknown until now. Recently, it was hypothesized that blood-seeking female tabanids prefer elevated temperatures, because their wing muscles are quicker and their nervous system functions better at a warmer body temperature brought about by warmer microclimate, and thus they can more successfully avoid the host's parasite-repelling reactions by prompt takeoffs. To test this hypothesis, we studied in field experiments the success rate of escape reactions of tabanids that landed on black targets as a function of the target temperature, and measured the surface temperature of differently coloured horses with thermography. We found that the escape success of tabanids decreased with decreasing target temperature, that is escape success is driven by temperature. Our results explain the behaviour of biting horseflies that they prefer warmer hosts against colder ones. Since in sunshine the darker the host the warmer its body surface, our results also explain why horseflies prefer sunlit dark (brown, black) hosts against bright (beige, white) ones, and why these parasites attack their hosts usually in sunshine, rather than under shaded conditions.

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.

Attractiveness of thermally different, uniformly black targets to horseflies: Tabanus tergestinus prefers sunlit warm shiny dark targets.

From a large distance tabanid flies may find their host animal by means of its shape, size, motion, odour, radiance and degree of polarization of host-reflected light. After alighting on the host, tabanids may use their mechano-, thermo-, hygro- and chemoreceptors to sense the substrate characteristics. Female tabanids prefer to attack sunlit against shady dark host animals, or dark against bright hosts for a blood meal, the exact reasons for which are unknown. Since sunlit darker surfaces are warmer than shady ones or sunlit/shady brighter surfaces, the differences in surface temperatures of dark and bright as well as sunlit and shady hosts may partly explain their different attractiveness to tabanids. We tested this observed warmth preference in field experiments, where we compared the attractiveness to tabanids (Tabanus tergestinus) of a warm and a cold shiny black barrel imitating dark hosts with the same optical characteristics. Using imaging polarimetry, thermography and Schlieren imaging, we measured the optical and thermal characteristics of both barrels and their small-scale models. We recorded the number of landings on these targets and measured the time periods spent on them. Our study revealed that T. tergestinus tabanid flies prefer sunlit warm shiny black targets against sunlit or shady cold ones with the same optical characteristics. These results support our new hypothesis that a blood-seeking female tabanid prefers elevated temperatures, partly because her wing muscles are more rapid and her nervous system functions better (due to faster conduction velocities and synaptic transmission of signals) in a warmer microclimate, and thus, she can avoid the parasite-repelling reactions of host animals by a prompt take-off.

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.

Experimental evidence that stripes do not cool zebras.

There are as many as 18 theories for the possible functions of the stripes of zebras, one of which is to cool the animal. We performed field experiments and thermographic measurements to investigate whether thermoregulation might work for zebra-striped bodies. A zebra body was modelled by water-filled metal barrels covered with horse, cattle and zebra hides and with various black, white, grey and striped patterns. The barrels were installed in the open air for four months while their core temperature was measured continuously. Using thermography, the temperature distributions of the barrel surfaces were compared to those of living zebras. The sunlit zebra-striped barrels reproduced well the surface temperature characteristics of sunlit zebras. We found that there were no significant core temperature differences between the striped and grey barrels, even on many hot days, independent of the air temperature and wind speed. The average core temperature of the barrels increased as follows: white cattle, grey cattle, real zebra, artificial zebra, grey horse, black cattle. Consequently, we demonstrate that zebra-striped coats do not keep the body cooler than grey coats challenging the hypothesis of a thermoregulatory role of zebra stripes.

Why do horseflies need polarization vision for host detection? Polarization helps tabanid flies to select sunlit dark host animals from the dark patches of the visual environment.

Horseflies (Tabanidae) are polarotactic, being attracted to linearly polarized light when searching for water or host animals. Although it is well known that horseflies prefer sunlit dark and strongly polarizing hosts, the reason for this preference is unknown. According to our hypothesis, horseflies use their polarization sensitivity to look for targets with higher degrees of polarization in their optical environment, which as a result facilitates detection of sunlit dark host animals. In this work, we tested this hypothesis. Using imaging polarimetry, we measured the reflection-polarization patterns of a dark host model and a living black cow under various illumination conditions and with different vegetation backgrounds. We focused on the intensity and degree of polarization of light originating from dark patches of vegetation and the dark model/cow. We compared the chances of successful host selection based on either intensity or degree of polarization of the target and the combination of these two parameters. We show that the use of polarization information considerably increases the effectiveness of visual detection of dark host animals even in front of sunny-shady-patchy vegetation. Differentiation between a weakly polarizing, shady (dark) vegetation region and a sunlit, highly polarizing dark host animal increases the efficiency of host search by horseflies.

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.