Graham R. Martin

Sexually selected traits such as feather ornamentation of male birds can act as an impediment to movement and predator detection. Here, we report a previously undocumented example of an impediment derived from a sexually selected trait: the cranial feather ornamentation in male Chrysolophus pheasants restricting their visual field. Visual fields define the space around an animal from which visual information can be retrieved. Out of the 300 bird species studied to date, there have been no significant differences reported in the visual fields between sexes. Our findings reveal that the cranial feathers of male golden ( C. pictus ) and Lady Amherst’s ( C. amherstiae ) pheasants significantly restrict their visual field relative to females and may impede their ability to gather information from the world about them. This effect is most extreme in the vertical extent, where a 30° and 40° difference is evident between the sexes of golden and Lady Amherst’s pheasants, respectively. The two Chrysolophus pheasant species are the first species studied to show a difference in visual fields between sexes; this difference was absent in two closely related species also measured in this study, silver pheasants ( Lophura nycthemera ) and green pheasants ( Phasianus versicolor ).

Wide variation in visual field configuration has been recorded among avian species and it is hypothesized that this variation is driven primarily by foraging ecology and predator detection. It has also been shown that visual field configurations can render some species more vulnerable to collisions with human artefacts that extend into open airspace, such as power lines and wind turbines. Visual fields have three main components: the monocular fields describe the extent of the world seen by each eye, the binocular field describes the region where the monocular fields overlap, and the blind area describes the region in which no vision is provided. Among birds, the topography of the binocular field, and the extent and position of the blind area, show considerable interspecific variation. Although Laridae (gulls, terns, skimmers) are a large and cosmopolitan taxon, visual field characteristics of only one species, Black Skimmer Rynchops niger , have been determined. However, skimmers are distinct from other Laridae species because they use a specialized foraging technique based upon tactile cues. We determined visual fields in three species of gulls (European Herring Gulls Larus argentatus , Lesser Black‐backed Gulls Larus fuscus , Black‐legged Kittiwakes Rissa tridactyla ), and found that they show the key characteristics associated with visually guided foraging. However, the binocular field does not extend through the full height of the frontal field. This results in a blind sector, which can project in the direction of flight when gulls pitch their heads sufficiently far forwards to visually search the surface below. This could render gulls vulnerable to collisions with anthropogenic structures (power lines, wind turbines) that extend into the open airspace. Photographs show that gulls in level flight do pitch their heads forward sufficiently to render them almost blind in the direction of travel, and further work on the head positions adopted by gulls in flight are recommended. The visual field of skimmers differs markedly from those of gulls. Their binocular field topography is interpreted as functioning in the control of bill position when skimming (flying just above the water surface with the elongated, blade‐like, rhamphotheca of the mandible extending through the water surface). Skimmers also have a blind area, which projects forwards in the direction of travel when skimming. This can be associated with the vulnerability of skimmers to collisions with objects that extend just above the water surface.

The binocular field of vision differs widely in birds depending on ecological traits such as foraging. Owls (Strigiformes) have been considered to have a unique binocular field, but whether it is related to foraging has remained unknown. While taking into account allometry and phylogeny, we hypothesized that both daily activity cycle and diet determine the size and shape of the binocular field in owls. Here, we compared the binocular field configuration of 23 species of owls. While we found no effect of allometry and phylogeny, ecological traits strongly influence the binocular field shape and size. Binocular field shape of owls significantly differed from that of diurnal raptors. Among owls, binocular field shape was relatively conserved, but binocular field size differed among species depending on ecological traits, with larger binocular fields in species living in dense habitat and foraging on invertebrates. Our results suggest that (i) binocular field shape is associated with the time of foraging in the daily cycle (owls versus diurnal raptors) and (ii) that binocular field size differs between closely related owl species even though the general shape is conserved, possibly because the field of view is partially restricted by feathers, in a trade-off with auditory localization.

The design of bird diverters should be based upon the perception of birds, not the perception of humans, but until now it is human vision that has guided diverter design. Aspects of bird vision pertinent to diverter design are reviewed. These are applied in an example that uses Canada Geese Branta canadensis as a putative worst-case example of a collision-prone species. The proposed design uses an achromatic checkerboard pattern of high contrast whose elements match the low spatial resolution of these birds when they are active under twilight light levels. The detectability of the device will be increased by movement, and this is best achieved with a device that rotates on its own axis driven by the wind. The recommended spacing of diverters along a power line is based upon the maximum width of the bird’s binocular field and the linear distance that it subtends at a distance sufficient to allow a bird to alter its flight path before possible impact. Given the worst-case nature of this example, other bird species should detect and avoid such a device. The basic design can be modified for use with specific target species if sufficient is known about their vision. Field trials of devices based on these design criteria are now required.

Although flight is regarded as a key behaviour of birds the perceptual demands for its control are met within constraints set by the perceptual demands of two other key tasks: the control of bill position, and the detection of food items/predators. Control of bill position, and timing its arrival at a target, are based upon information derived from the optic flow-field in the binocular region that encompasses the bill. It uses information extracted from close to the bird using vision of relatively low spatial resolution. The detection of food items and predators is based upon information detected at a greater distance and depends upon regions in the retina with relatively high spatial resolution. The tasks of detecting predators and of placing the bill accurately, make contradictory demands upon vision and these have resulted in trade-offs in the form of visual fields and in the topography of retinal regions in which spatial resolution is enhanced, indicated by foveas, areas and high ganglion cell densities. The informational function of binocular vision in birds does not lie in binocularity per se (i.e. two eyes receiving slightly different information simultaneously about the same objects) but in the contralateral projection of the visual field of each eye. This ensures that each eye receives information from a symmetrically expanding optic flow-field centred close to the direction of the bill, and from this the crucial information of direction of travel and time-to-contact can be extracted, almost instantaneously. Interspecific comparisons of visual fields between closely related species have shown that small differences in foraging techniques can give rise to different perceptual challenges and these have resulted in differences in visual fields even within the same genus. This suggests that vision is subject to continuing and relatively rapid natural selection based upon individual differences in the structure of the optical system, retinal topography and eye position in the skull. From a sensory ecology perspective a bird is best characterised as “a bill guided by an eye” and that control of flight is achieved within constraints on visual capacity dictated primarily by the demands of foraging and bill control.

Significant differences in avian visual fields are found between closely related species that differ in their foraging technique. We report marked differences in the visual fields of two auk species. In air, Common Guillemots Uria aalge have relatively narrow binocular fields typical of those found in non‐passerine predatory birds. Atlantic Puffins Fratercula arctica have much broader binocular fields similar to those that have hitherto been recorded in passerines and in a penguin. In water, visual fields narrow considerably and binocularity in the direction of the bill is probably abolished in both auk species. Although perceptual challenges associated with foraging are similar in both species during the breeding season, when they are piscivorous, Puffins (but not Guillemots) face more exacting perceptual challenges when foraging at other times, when they take a high proportion of small invertebrate prey. Capturing this prey probably requires more accurate, visually guided bill placement and we argue that this is met by the Puffin's broader binocular field, which is retained upon immersion; its upward orientation may enable prey to be seen in silhouette. These visual field configurations have potentially important consequences that render these birds vulnerable to collision with human artefacts underwater, but not in air. They also have consequences for vigilance behaviour.