Many animals have the potential to discriminate nonspectral colors. For humans, purple is the clearest example of a nonspectral color. It is perceived when two color cone types in the retina (blue and red) with nonadjacent spectral sensitivity curves are predominantly stimulated. Purple is considered nonspectral because no monochromatic light (such as from a rainbow) can evoke this simultaneous stimulation. Except in primates and bees, few behavioral experiments have directly examined nonspectral color discrimination, and little is known about nonspectral color perception in animals with more than three types of color photoreceptors. Birds have four color cone types (compared to three in humans) and might perceive additional nonspectral colors such as UV+red and UV+green. Can birds discriminate nonspectral colors, and are these colors behaviorally and ecologically relevant? Here, using comprehensive behavioral experiments, we show that wild hummingbirds can discriminate a variety of nonspectral colors. We also show that hummingbirds, relative to humans, likely perceive a greater proportion of natural colors as nonspectral. Our analysis of plumage and plant spectra reveals many colors that would be perceived as nonspectral by birds but not by humans: Birds’ extra cone type allows them not just to see UV light but also to discriminate additional nonspectral colors. Our results support the idea that birds can distinguish colors throughout tetrachromatic color space and indicate that nonspectral color perception is vital for signaling and foraging. Since tetrachromacy appears to have evolved early in vertebrates, this capacity for rich nonspectral color perception is likely widespread.

Lay SummaryThe influence of coloration on the ecology of moving animals in groups is not well understood. High-contrast stripes may act as “motion dazzle camouflage” by preventing predators from accurately tracking prey, particularly in groups. We used a computer study with human “predators” to investigate this and found that targets in groups with horizontal striped patterns were harder to track than the ones with conventional camouflage, and the benefit increased with group size.Twitter: @Camo_Lab

‘Motion dazzle camouflage’ is the name for the putative effects of highly conspicuous, often repetitive or complex, patterns on parameters important in prey capture, such as the perception of speed, direction and identity. Research into motion dazzle camouflage is increasing our understanding of the interactions between visual tracking, the confusion effect and defensive coloration. However, there is a paucity of research into the effects of contrast on motion dazzle camouflage: is maximal contrast a prerequisite for effectiveness? If not, this has important implications for our recognition of the phenotype and understanding of the function and mechanisms of potential motion dazzle camouflage patterns. Here we tested human participants' ability to track one moving target among many identical distractors with surface patterns designed to test the influence of these factors. In line with previous evidence, we found that targets with stripes parallel to the object direction of motion were hardest to track. However, reduction in contrast did not significantly influence this result. This finding may bring into question the utility of current definitions of motion dazzle camouflage, and means that some animal patterns, such as aposematic or mimetic stripes, may have previously unrecognized multiple functions.

Many animal signals are complex, often combining multimodal components with dynamic motion. To understand the function and evolution of these displays, it is vital to appreciate their spatiotemporal organization. Male broad-tailed hummingbirds (Selasphorus platycercus) perform dramatic U-shaped courtship dives over females, appearing to combine rapid movement and dive-specific mechanical noises with visual signals from their iridescent gorgets. To understand how motion, sound and color interact in these spectacular displays, we obtained video and audio recordings of dives performed by wild hummingbirds. We then applied a multi-angle imaging technique to estimate how a female would perceive the male’s iridescent gorget throughout the dive. We show that the key physical, acoustic and visual aspects of the dive are remarkably synchronized—all occurring within 300 milliseconds. Our results highlight the critical importance of accounting for motion and orientation when investigating animal displays: speed and trajectory affect how multisensory signals are produced and perceived.

One contribution of 18 to a theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.Despite a recent explosion of research on pattern recognition, in both neuroscience and computer vision, we lack a basic understanding of how most animals perceive and respond to patterns in the wild. Avian brood parasites and their hosts provide an ideal study system for investigating the mechanisms of pattern recognition. The cuckoo finch, Anomalospiza imberbis, and its host the tawny-flanked prinia, Prinia subflava, lay highly polymorphic eggs with a great deal of variation in colour and patterning, with the cuckoo finch capable of close egg mimicry. Behavioural experiments in Zambia have previously shown that prinias use colour and multiple 'low-level' (occurring in early stages of visual processing) pattern attributes, derived from spatial frequency analysis, when rejecting foreign eggs. Here, we explore the extent to which host birds might also use 'higher-level' pattern attributes, derived from a feature detection algorithm, to make rejection decisions. Using a SIFT-based pattern recognition algorithm, NATUREPATTERNMATCH, we show that hosts are more likely to reject a foreign egg if its higher-level pattern features-which capture information about the shape and orientation of markings-differ from those of the host eggs. A revised statistical model explains about 37% variance in egg rejection behaviour, and differences in colour, low-level and higher-level pattern features all predict rejection, accounting for 42, 44 and 14% of the explained variance, respectively. Thus, higher-level pattern features provide a small but measurable improvement to the original model and may be especially useful when colour and low-level pattern features provide hosts with little information. Understanding the relative importance of low-and higher-level pattern features is a valuable goal for future work on animal coloration, especially in the contexts of mimicry, camouflage and individual recognition.This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

The confusion effect describes the phenomenon of decreasing predator attack success with increasing prey group size. However, there is a paucity of research into the influence of this effect in coherent groups, such as flocks of European starlings (Sturnus vulgaris). Here, for the first time, we use a computer game style experiment to investigate the confusion effect in three dimensions. To date, computerized studies on the confusion effect have used two-dimensional simulations with simplistic prey movement and dynamics. Our experiment is the first investigation of the effects of flock size and density on the ability of a (human) predator to track and capture a target starling in a realistically simulated three-dimensional flock of starlings. In line with the predictions of the confusion effect, modelled starlings appear to be safer from predation in larger and denser flocks. This finding lends credence to previous suggestions that starling flocks have anti-predator benefits and, more generally, it suggests that active increases in density in animal groups in response to predation may increase the effectiveness of the confusion effect.

Brood parasites use the parental care of others to raise their young and sometimes employ mimicry to dupe their hosts. The brood-parasitic finches of the genus Vidua are a textbook example of the role of imprinting in sympatric speciation. Sympatric speciation is thought to occur in Vidua because their mating traits and host preferences are strongly influenced by their early host environment. However, this alone may not be sufficient to isolate parasite lineages, and divergent ecological adaptations may also be required to prevent hybridization collapsing incipient species. Using pattern recognition software and classification models, we provide quantitative evidence that Vidua exhibit specialist mimicry of their grassfinch hosts, matching the patterns, colors and sounds of their respective host's nestlings. We also provide qualitative evidence of mimicry in postural components of Vidua begging. Quantitative comparisons reveal small discrepancies between parasite and host phenotypes, with parasites sometimes exaggerating their host's traits. Our results support the hypothesis that behavioral imprinting on hosts has not only enabled the origin of new Vidua species, but also set the stage for the evolution of host-specific, ecological adaptations.

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