A hypothesis for robust polarization vision: an example from the Australian imperial blue butterfly, Jalmenus evagoras.

The Australian lycaenid butterfly Jalmenus evagoras has iridescent wings that are sexually dimorphic, spectrally and in their degree of polarization, suggesting that these properties are likely to be important in mate recognition. We first describe the results of a field experiment showing that free-flying individuals of J. evagoras discriminate between visual stimuli that vary in polarization content in blue wavelengths but not in others. We then present detailed reflectance spectrophotometry measurements of the polarization content of male and female wings, showing that female wings exhibit blue-shifted reflectance, with a lower degree of polarization relative to male wings. Finally, we describe a novel method for measuring alignment of ommatidial arrays: by measuring variation of depolarized eyeshine intensity from patches of ommatidia as a function of eye rotation, we show that (a) individual rhabdoms contain mutually perpendicular microvilli; (b) many rhabdoms in the array have their microvilli misaligned with respect to neighboring rhabdoms by as much as 45 deg; and (c) the misaligned ommatidia are useful for robust polarization detection. By mapping the distribution of the ommatidial misalignments in eye patches of J. evagoras, we show that males and females exhibit differences in the extent to which ommatidia are aligned. Both the number of misaligned ommatidia suitable for robust polarization detection and the number of aligned ommatidia suitable for edge detection vary with respect to both sex and eye patch elevation. Thus, J. evagoras exhibits finely tuned ommatidial arrays suitable for perception of polarized signals, likely to match sex-specific life history differences in the utility of polarized signals.

Egg patterns as identity signals in colonial seabirds: a comparison of four alcid species.

The ability to recognize mates, kin, offspring and neighbors by their individually distinctive traits-individual recognition (IR)-is widespread in animals. Much work has investigated IR from the perspective of the recognizer, but less is known about the extent to which signals have evolved to facilitate IR. To explore this, one approach is to compare putative identity signals among species that differ in life history and extent of IR. In Common Murres (Uria aalge), a colonially breeding seabird, the eggs of individual females are remarkably variable in terms of color and pattern (maculation). Common Murres also appear to recognize their own eggs, leading to the hypothesis that variable egg phenotypes evolved to promote recognizability. However, we lack a quantitative assessment of the egg pattern information in Common Murres and their close relatives. Here, we analyzed images of eggs laid by four alcid species: Common Murres, Thick-billed Murres (Uria lomvia), Razorbills (Alca torda) and Dovekies (Alle alle). We extracted pattern measures believed to be relevant to bird vision and calculated Beecher's information statistic (Hs ), which allowed us to compare the amount of identity information contained in each species' egg patterns. Murres, which nest in dense colonies and can recognize their own eggs, have egg patterns with a relatively large amount of identity information compared to Razorbills and Dovekies. Egg recognition has not been demonstrated in Razorbills and Dovekies, whose colonies are less dense. Our results are consistent with the hypothesis that complex patterns of Murre eggs may have evolved to increase individual recognizability.

Multimodal mimicry of hosts in a radiation of parasitic finches.

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.

Wild hummingbirds discriminate nonspectral colors.

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.

Higher-level pattern features provide additional information to birds when recognizing and rejecting parasitic eggs.

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'.

Synchronization of speed, sound and iridescent color in a hummingbird aerial courtship dive.

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.

The confusion effect when attacking simulated three-dimensional starling flocks.

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.

Dazzle camouflage and the confusion effect: the influence of varying speed on target tracking.

The formation of groups is a common strategy to avoid predation in animals, and recent research has indicated that there may be interactions between some forms of defensive coloration, notably high-contrast 'dazzle camouflage', and one of the proposed benefits of grouping: the confusion effect. However, research into the benefits of dazzle camouflage has largely used targets moving with constant speed. This simplification may not generalize well to real animal systems, where a number of factors influence both within- and between-individual variation in speed. Departure from the speed of your neighbours in a group may be predicted to undermine the confusion effect. This is because individual speed may become a parameter through which the observer can individuate otherwise similar targets: an 'oddity effect'. However, dazzle camouflage patterns are thought to interfere with predator perception of speed and trajectory. The current experiment investigated the possibility that such patterns could ameliorate the oddity effect caused by within-group differences in prey speed. We found that variation in speed increased the ease with which participants could track targets in all conditions. However, we found no evidence that motion dazzle camouflage patterns reduced oddity effects based on this variation in speed, a result that may be informative about the mechanisms behind this form of defensive coloration. In addition, results from those conditions most similar to those of published studies replicated previous results, indicating that targets with stripes parallel to the direction of motion are harder to track, and that this pattern interacts with the confusion effect to a greater degree than background matching or orthogonal-to-motion striped patterns.

Dazzle camouflage, target tracking, and the confusion effect.

The influence of coloration on the ecology and evolution of moving animals in groups is poorly understood. Animals in groups benefit from the "confusion effect," where predator attack success is reduced with increasing group size or density. This is thought to be due to a sensory bottleneck: an increase in the difficulty of tracking one object among many. Motion dazzle camouflage has been hypothesized to disrupt accurate perception of the trajectory or speed of an object or animal. The current study investigates the suggestion that dazzle camouflage may enhance the confusion effect. Utilizing a computer game style experiment with human predators, we found that when moving in groups, targets with stripes parallel to the targets' direction of motion interact with the confusion effect to a greater degree, and are harder to track, than those with more conventional background matching patterns. The findings represent empirical evidence that some high-contrast patterns may benefit animals in groups. The results also highlight the possibility that orientation and turning may be more relevant in the mechanisms of dazzle camouflage than previously recognized.

Contrast, contours and the confusion effect in dazzle camouflage.

'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.