Antipredator defences in motion: animals reduce predation risks by concealing or misleading motion signals.

Motion is a crucial part of the natural world, yet our understanding of how animals avoid predation whilst moving remains rather limited. Although several theories have been proposed for how antipredator defence may be facilitated during motion, there is often a lack of supporting empirical evidence, or conflicting findings. Furthermore, many studies have shown that motion often 'breaks' camouflage, as sudden movement can be detected even before an individual is recognised. Whilst some static camouflage strategies may conceal moving animals to a certain extent, more emphasis should be given to other modes of camouflage and related defences in the context of motion (e.g. flicker fusion camouflage, active motion camouflage, motion dazzle, and protean motion). Furthermore, when motion is involved, defence strategies are not necessarily limited to concealment. An animal can also rely on motion to mislead predators with regards to its trajectory, location, size, colour pattern, or even identity. In this review, we discuss the various underlying antipredator strategies and the mechanisms through which they may be linked to motion, conceptualising existing empirical and theoretical studies from two perspectives - concealing and misleading effects. We also highlight gaps in our understanding of these antipredator strategies, and suggest possible methodologies for experimental designs/test subjects (i.e. prey and/or predators) and future research directions.

Background matching can reduce responsiveness of jumping spiders to stimuli in motion.

Motion and camouflage were previously considered to be mutually exclusive, as sudden movements can be easily detected. Background matching, for instance, is a well-known, effective camouflage strategy where the colour and pattern of a stationary animal match its surrounding background. However, background matching may lose its efficacy when the animal moves, as the boundaries of the animal become more defined against its background. Recent evidence shows otherwise, as camouflaged objects can be less detectable than uncamouflaged objects even while in motion. Here, we explored whether the detectability of computer-generated stimuli varies with the speed of motion, background (matching and unmatching) and size of stimuli in six species of jumping spiders (Araneae: Salticidae). Our results showed that, in general, the responsiveness of all six salticid species tested decreased with increasing stimulus speed regardless of whether the stimuli were conspicuous or camouflaged. Importantly, salticid responses to camouflaged stimuli were significantly lower compared with those to conspicuous stimuli. There were significant differences in motion detectability across species when the stimuli were conspicuous, suggesting differences in visual acuity in closely related species of jumping spiders. Furthermore, small stimuli elicited significantly lower responses than large stimuli across species and speeds. Our results thus suggest that background matching is effective even when stimuli are in motion, reducing the detectability of moving stimuli.

Motion: enhancing signals and concealing cues.

Animal colour patterns remain a lively focus of evolutionary and behavioural ecology, despite the considerable conceptual and technical developments over the last four decades. Nevertheless, our current understanding of the function and efficacy of animal colour patterns remains largely shaped by a focus on stationary animals, typically in a static background. Yet, this rarely reflects the natural world: most animals are mobile in their search for food and mates, and their surrounding environment is usually dynamic. Thus, visual signalling involves not only animal colour patterns, but also the patterns of animal motion and behaviour, often in the context of a potentially dynamic background. While motion can reveal information about the signaller by attracting attention or revealing signaller attributes, motion can also be a means of concealing cues, by reducing the likelihood of detection (motion camouflage, motion masquerade and flicker-fusion effect) or the likelihood of capture following detection (motion dazzle and confusion effect). The interaction between the colour patterns of the animal and its local environment is further affected by the behaviour of the individual. Our review details how motion is intricately linked to signalling and suggests some avenues for future research. This Review has an associated Future Leader to Watch interview with the first author.

What's in a band? The function of the color and banding pattern of the Banded Swallowtail.

Butterflies have evolved a diversity of color patterns, but the ecological functions for most of these patterns are still poorly understood. The Banded Swallowtail butterfly, Papilio demolion demolion, is a mostly black butterfly with a greenish-blue band that traverses the wings. The function of this wing pattern remains unknown. Here, we examined the morphology of black and green-blue colored scales, and how the color and banding pattern affects predation risk in the wild. The protective benefits of the transversal band and of its green-blue color were tested via the use of paper model replicas of the Banded Swallowtail with variations in band shape and band color in a full factorial design. A variant model where the continuous transversal green-blue band was shifted and made discontinuous tested the protective benefit of the transversal band, while grayscale variants of the wildtype and distorted band models assessed the protective benefit of the green-blue color. Paper models of the variants and the wildtype were placed simultaneously in the field with live baits. Wildtype models were the least preyed upon compared with all other variants, while gray models with distorted bands suffered the greatest predation. The color and the continuous band of the Banded Swallowtail hence confer antipredator qualities. We propose that the shape of the band hinders detection of the butterfly's true shape through coincident disruptive coloration; while the green color of the band prevents detection of the butterfly from its background via differential blending. Differential blending is aided by the green-blue color being due to pigments rather than via structural coloration. Both green and black scales have identical structures, and the scales follow the Bauplan of pigmented scales documented in other Papilio butterflies.

Detritus decorations of an orb-weaving spider, Cyclosa mulmeinensis (Thorell): for food or camouflage?

Many species of the orb-web spider genus Cyclosa often adorn their webs with decorations of prey remains, egg sacs and/or plant detritus, termed ;detritus decorations'. These detritus decorations have been hypothesised to camouflage the spider from predators or prey and thus reduce predation risk or increase foraging success. In the present study, we tested these two alternative hypotheses simultaneously using two types of detritus decorations (prey remain and egg sac) built by Cyclosa mulmeinensis (Thorell). By monitoring the possible responses of predators to spiders on their webs with and without decorations in the field, we tested whether web decorations would reduce the mortality of spiders. Wasp predators were observed to fly in the vicinity of webs with decorations slightly more often than in the vicinity of webs without decorations but there were very few attacks on spiders by wasps. By comparing the insect interception rates of webs with and without decorations in the field, we tested whether web decorations would increase the foraging success. Webs decorated with prey remains or egg sacs intercepted more insects than those without in the field. By calculating colour contrasts of both prey-remain and egg-sac decorations against spiders viewed by bird (blue tits) and hymenopteran (e.g. wasps) predators as well as hymenopteran (bees) prey, we showed that C. mulmeinensis spiders on webs with egg-sac decorations were invisible to both hymenopteran prey and predators and bird predators over short and long distances. While spiders on webs with prey-remain decorations were invisible to both hymenopterans and birds over short distances, spiders on webs with prey-remain decorations were visible to both predators and prey over long distances. Our results thus suggest that decorating webs with prey remains and egg sacs in C. mulmeinensis may primarily function as camouflage to conceal the spider from insects rather than as prey attractants, possibly contributing to the interception of more insect prey. However, the detritus decorations exhibit varying success as camouflage against predators, depending on whether predators are jumping spiders, wasps or birds, as well as on the decoration type.