Antipredatory function of head shape for vipers and their mimics.

Most research into the adaptive significance of warning signals has focused on the colouration and patterns of prey animals. However, behaviour, odour and body shape can also have signal functions and thereby reduce predators' willingness to attack defended prey. European vipers all have a distinctive triangular head shape; and they are all venomous. Several non-venomous snakes, including the subfamily Natricinae, commonly flatten their heads (also known as head triangulation) when disturbed. The adaptive significance of this potential behavioural mimicry has never been investigated. We experimentally tested if the triangular head shape typical of vipers offers protection against predation.…

From deception to frankness: Benefits of ontogenetic shift in the anti-predator strategy of alder moth Acronicta alni larvae

Abstract Animals can avoid prédation by masquerading as objects that are not food to their predators. Alder moth Acronicta alni larvae go through an impressive ontogenetic change from masquerade to highly conspicuous appearance: early larval stages resemble bird droppings but in the last instar the larval coloration changes into striking yellow-and-black stripes. We hypothesized that such a change may be driven by differential prédation favoring dissimilar anti-predator strategies in different life stages. We show with a series of laboratory assays that larvae are distasteful to birds regardless of their developmental stage, suggesting that ontogenetic color change is not driven by the diff…

A case study of the coconut crab Birgus latro on Zanzibar highlights global threats and conservation solutions

Abstract The coconut crab Birgus latro, the largest terrestrial decapod, is under threat in most parts of its geographical range. Its life cycle involves two biomes (restricted terrestrial habitats near the coast, and salt water currents of the tropical Indian and Pacific Oceans). Its dependence on coastal habitat means it is highly vulnerable to the habitat destruction that typically accompanies human population expansion along coastlines. Additionally, it has a slow reproductive rate and can reach large adult body sizes that, together with its slow movement when on land, make it highly susceptible to overharvesting. We studied the distribution and population changes of coconut crabs at 15…

Temporal relationship between genetic and warning signal variation in the aposematic wood tiger moth (Parasemia plantaginis).

Many plants and animals advertise unpalatability through warning signals in the form of colour and shape. Variation in warning signals within local populations is not expected because they are subject to directional selection. However, mounting evidence of warning signal variation within local populations suggests that other selective forces may be acting. Moreover, different selective pressures may act on the individual components of a warning signal. At present, we have a limited understanding about how multiple selection processes operate simultaneously on warning signal components, and even less about their temporal and spatial dynamics. Here, we examined temporal variation of several w…

Context-dependent coloration of prey and predator decision making in contrasting light environments

A big question in behavioral ecology is what drives diversity of color signals. One possible explanation is that environmental conditions, such as light environment, may alter visual signaling of prey, which could affect predator decision-making. Here, we tested the context-dependent predator selection on prey coloration. In the first experiment, we tested detectability of artificial visual stimuli to blue tits (Cyanistes caeruleus) by manipulating stimulus luminance and chromatic context of the background. We expected the presence of the chromatic context to facilitate faster target detection. As expected, blue tits found targets on chromatic yellow background faster than on achromatic gre…

Geographic mosaic of selection by avian predators on hindwing warning colour in a polymorphic aposematic moth

AbstractWarning signals are predicted to develop signal monomorphism via positive frequency-dependent selection (+FDS) albeit many aposematic systems exhibit signal polymorphism. To understand this mismatch, we conducted a large-scale predation experiment in four locations, among which the frequencies of hindwing warning coloration of aposematic Arctia plantaginis differ. Here we show that selection by avian predators on warning colour is predicted by local morph frequency and predator community composition. We found +FDS to be strongest in monomorphic Scotland, and in contrast, lowest in polymorphic Finland, where different predators favour different male morphs. +FDS was also found in Geo…

Adaptation of a tropical butterfly to a temperate climate

Developmental plasticity enables organisms to cope with environmental heterogeneity, such as seasonal variation in climatic conditions, and is thought to affect a species’ capability to adapt to environments with novel seasonal and ecological dynamics. We studied developmental plasticity of the widespread tropical butterfly, Bicyclus safitza, which reaches the southern edge of its distribution in the temperate zone of South Africa. In wet–dry seasonal environments in tropical Africa, adults of Bicyclus butterflies are present all year round and exhibit discrete seasonal forms in alternating generations. We demonstrate that a population that colonized a more temperate climate region has adop…

The Eurasian Treecreeper (Certhia familiaris) has an effective camouflage against mammalian but not avian vision in boreal forests

A well-known example of visual camouflage in birds is the plumage coloration of the Eurasian Treecreeper Certhia familiaris, yet this species’ camouflage has never been objectively quantified. Here, we quantify treecreeper camouflage in its boreal forest habitat, test whether treecreepers better match tree backgrounds at nest site, territory or habitat spatial scales, and explore which common tree species provide the best background match. Using photographic data of the birds and forested backgrounds, we test their background match using human, ferret and avian vision modelling. We found that a treecreeper’s wing and mantle provided closest background matching, whereas the wing stripe and t…

Many forms of the wood tiger moth (Parasemia plantaginis) : selective heterogeneity favours polymorphic warning signals

Camouflage in arid environments: the case of Sahara-Sahel desert rodents

Deserts and semi-deserts, such as the Sahara-Sahel region in North Africa, are exposed environments with restricted vegetation coverage. Due to limited physical surface structures, these open areas provide a promising ecosystem to understand selection for crypsis. Here, we review knowledge on camouflage adaptation in the Sahara-Sahel rodent community, which represents one of the best documented cases of phenotype-environment convergence comprising a marked taxonomic diversity. Through their evolutionary history, several rodent species from the Sahara-Sahel have repeatedly evolved an accurate background matching against visually-guided predators. Top-down selection by predators is therefore …

The effect of spatial and temporal scale on camouflage in North African rodents

Abstract Background matching, a common form of camouflage, is a widespread anti-predator adaptation that hinders detection or recognition by increasing the resemblance of prey to its environment. However, the natural environment is complex and both spatially and temporally variable, which constrains effective background matching as an anti-predator strategy. Here, using remote sensing data (publicly available satellite imagery), we investigated how variation of habitat parameters predicts background matching in 16 Sahara–Sahel rodent species across spatial and temporal scales. All fur colour parameters (hue, saturation and brightness) strongly matched the respective habitats of the differen…

Transparency reduces predator detection in chemically protected clearwing butterflies

Abstract1. Predation is an important selective pressure and some prey have evolved warning colour signals advertising unpalatability (i.e. aposematism) as an antipredator strategy. Unexpectedly, some butterfly species from the unpalatable tribe Ithomiini possess transparent wings, an adaptation rare on land but common in water where it helps avoiding predator detection.2. We tested if transparency of butterfly wings was associated with decreased detectability by predators, by comparing four butterfly species exhibiting different degrees of transparency, ranging from fully opaque to largely transparent. We tested our prediction using using both wild birds and humans in behavioural experiment…

Trade-off between warning signal efficacy and mating success in the wood tiger moth

The coloration of species can have multiple functions, such as predator avoidance and sexual signalling, that directly affect fitness. As selection should favour traits that positively affect fitness, the genes underlying the trait should reach fixation, thereby preventing the evolution of polymorphisms. This is particularly true for aposematic species that rely on coloration as a warning signal to advertise their unprofitability to predators. Nonetheless, there are numerous examples of aposematic species showing remarkable colour polymorphisms. We examined whether colour polymorphism in the wood tiger moth is maintained by trade-offs between different functions of coloration. In Finland, m…

Multimodal Aposematic Signals and Their Emerging Role in Mate Attraction

Chemically defended animals often display conspicuous color patterns that predators learn to associate with their unprofitability and subsequently avoid. Such animals (i.e., aposematic), deter predators by stimulating their visual and chemical sensory channels. Hence, aposematism is considered to be “multimodal.” The evolution of warning signals (and to a lesser degree their accompanying chemical defenses) is fundamentally linked to natural selection by predators. Lately, however, increasing evidence also points to a role of sexual selection shaping warning signal evolution. One of the species in which this has been shown is the wood tiger moth, Arctia plantaginis, which we here put forward…

Adaptation of a tropical butterfly to a temperate climate

Bean goose migration shows a long-term temporal shift to earlier spring, but not to later autumn migration in Finland

Ilmastonmuutos vaatii eliöitä mukauttamaan perinnöllisiä tai opittuja käyttäytymismallejaan muuttuviin ympäristöolosuhteisiin. Erityisen alttiita muutoksille voi olla erilaisten habitaattien välillä muuttavat linnut, joiden selviytyminen on yhteydessä muuton onnistuneeseen ajoittamiseen. Tässä tutkimuksessa käytimme metsähanhesta (Anser fabalis) avoimeen lintutietopalveluun, Tiiraan (tiira.fi – BirdLife Suomi), kerättyjä havaintoja selvittääksemme, onko lajin muuton ajoituksessa tapahtunut muutosta viimeisten 40 vuoden aikana. Lisäksi selvitimme ajoituksen yhteyttä muuton aikaiseen suursäätilaan käyttämällä Pohjois-Atlantin suursäätilaa kuvaavaa NAO-indeksiä. Tutkitulla ajanjaksolla 1978–20…

Genetic colour variation visible for predators and conspecifics is concealed from humans in a polymorphic moth

The definition of colour polymorphism is intuitive: genetic variants express discretely coloured phenotypes. This classification is, however, elusive as humans form subjective categories or ignore differences that cannot be seen by human eyes. We demonstrate an example of a 'cryptic morph' in a polymorphic wood tiger moth (Arctia plantaginis), a phenomenon that may be common among well-studied species. We used pedigree data from nearly 20,000 individuals to infer the inheritance of hindwing colouration. The evidence supports a single Mendelian locus with two alleles in males: WW and Wy produce the white and yy the yellow hindwing colour. The inheritance could not be resolved in females as t…

The giant panda is cryptic

The giant panda (Ailuropoda melanoleuca) is an iconic mammal, but the function of its black-and-white coloration is mysterious. Using photographs of giant pandas taken in the wild and state-of-the-art image analysis, we confirm the counterintuitive hypothesis that their coloration provides camouflage in their natural environment. The black fur blends into dark shades and tree trunks, whereas white fur matches foliage and snow when present, and intermediate pelage tones match rocks and ground. At longer viewing distances giant pandas show high edge disruption that breaks up their outline, and up close they rely more on background matching. The results are consistent across acuity-corrected c…

Through predators’ eyes: phenotype–environment associations in shore crab coloration at different spatial scales

We thank Emil Aaltonen Foundation providing a young scientist grant for this research (O.N.). MS and AEL were supported by a Biotechnology and Biological Sciences Research Council (BBSRC) David Phillips Research Fellowship (BB/G022887/1).

Camouflage accuracy in Sahara-Sahel desert rodents

1.Camouflage helps animals to hide from predators and is therefore key to survival. Although widespread convergence of animal phenotypes to their natural environment is well established, there is a lack of knowledge about how species compromise camouflage accuracy across different background types in their habitat. 2.Here we tested how background matching has responded to top‐down selection by avian and mammalian predators using Sahara‐Sahel desert rodents in North Africa. 3.We show that the fur coloration of several species has become an accurate match to different types of desert habitats. This is supported by a correlation analysis of colour and pattern metrics, investigation of animal‐t…

Environment-mediated morph-linked immune and life-history responses in the aposematic wood tiger moth

1. Warning signals are expected to evolve towards conspicuousness and monomorphism, and thereby hamper the evolution of multiple colour morphs. Here, we test fitness responses to different rearing densities to explain colour polymorphism in aposematic wood tiger moth (Parasemia plantaginis) males. 2. We used larval lines sired by white or yellow adult males selected for small or large melanization patterns of coloration. We reared these selected lines either solitarily (favourable conditions) or in aggregations (challenged conditions), and followed their performance to adult stage. We tested whether differences in larval density affected life-history traits, adult melanin expression, adult …

Behavioural thresholds of blue tit colour vision and the effect of background chromatic complexity

Vision is a vital attribute to foraging, navigation, mate selection and social signalling in animals, which often have a very different colour perception in comparison to humans. For understanding how animal colour perception works, vision models provide the smallest colour difference that animals of a given species are assumed to detect. To determine the just-noticeable-difference, or JND, vision models use Weber fractions that set discrimination thresholds of a stimulus compared to its background. However, although vision models are widely used, they rely on assumptions of Weber fractions since the exact fractions are unknown for most species. Here, we test; i) which Weber fractions in lo…

Colour polymorphism torn apart by opposing positive frequency-dependent selection, yet maintained in space.

Summary Polymorphic warning signals in aposematic species are enigmatic because predator learning and discrimination should select for the most common coloration, resulting in positive frequency‐dependent survival selection. Here, we investigated whether differential mating success could create sufficiently strong negative frequency‐dependent selection for rare morphs to explain polymorphic (white and yellow) warning coloration in male wood tiger moths (Parasemia plantaginis). We conducted an experiment in semi‐natural conditions where we estimated mating success for both white and yellow male moths under three different morph frequencies. Contrary to expectations, mating success was positi…

Camouflage accuracy in Sahara–Sahel desert rodents

1. Camouflage helps animals to hide from predators and is therefore key to survival. Although widespread convergence of animal phenotypes to their natural environment is well-established, there is a lack of knowledge about how species compromise camouflage accuracy across different background types in their habitat.2. Here we tested how background matching has responded to top–down selection by avian and mammalian predators using Sahara–Sahel desert rodents in North Africa.3. We show that the fur colouration of several species has become an accurate match to different types of desert habitats. This is supported by a correlation analysis of colour and pattern metrics, investigation of animal…

To quiver or to shiver: increased melanization benefits thermoregulation, but reduces warning signal efficacy in the wood tiger moth

Melanin production is often considered costly, yet beneficial for thermoregulation. Studies of variation in melanization and the opposing selective forces that underlie its variability contribute greatly to understanding natural selection. We investigated whether melanization benefits are traded off with predation risk to promote observed local and geographical variation in the warning signal of adult male wood tiger moths ( Parasemia plantaginis ). Warning signal variation is predicted to reduce survival in aposematic species. However, in P. plantaginis , male hindwings are either yellow or white in Europe, and show continuous variation in melanized markings that cover 20 to 90 per cent o…

Variable crab camouflage patterns defeat search image formation.

Understanding what maintains the broad spectrum of variation in animal phenotypes and how this influences survival is a key question in biology. Frequency dependent selection – where predators temporarily focus on one morph at the expense of others by forming a “search image” – can help explain this phenomenon. However, past work has never tested real prey colour patterns, and rarely considered the role of different types of camouflage. Using a novel citizen science computer experiment that presented crab “prey” to humans against natural backgrounds in specific sequences, we were able to test a range of key hypotheses concerning the interactions between predator learning, camouflage and mor…

Multimodal Aposematic Defenses Through the Predation Sequence

Aposematic organisms warn predators of their unprofitability using a combination of defenses, including visual warning signals, startling sounds, noxious odors, or aversive tastes. Using multiple lines of defense can help prey avoid predators by stimulating multiple senses and/or by acting at different stages of predation. We tested the efficacy of three lines of defense (color, smell, taste) during the predation sequence of aposematic wood tiger moths (Arctia plantaginis) using blue tit (Cyanistes caeruleus) predators. Moths with two hindwing phenotypes (genotypes: WW/Wy = white, yy = yellow) were manipulated to have defense fluid with aversive smell (methoxypyrazines), body tissues with a…

Transparency reduces predator detection in mimetic clearwing butterflies

International audience; Predation is an important selective pressure and some prey have evolved conspicuous warning signals that advertise unpalatability (i.e. aposematism) as an antipredator defence. Conspicuous colour patterns have been shown effective as warning signals, by promoting predator learning and memory. Unexpectedly, some butterfly species from the unpalatable tribe Ithomiini possess transparent wings, a feature rare on land but common in water, known to reduce predator detection.We tested if transparency of butterfly wings was associated with decreased detectability by predators, by comparing four butterfly species exhibiting different degrees of transparency, ranging from ful…

Improved camouflage through ontogenetic colour change confers reduced detection risk in shore crabs

Abstract Animals from many taxa, from snakes and crabs to caterpillars and lobsters, change appearance with age, but the reasons why this occurs are rarely tested.We show the importance that ontogenetic changes in coloration have on the camouflage of the green shore crabs (Carcinus maenas), known for their remarkable phenotypic variation and plasticity in colour and pattern.In controlled conditions, we reared juvenile crabs of two shades, pale or dark, on two background types simulating different habitats for 10 weeks.In contrast to expectations for reversible colour change, crabs did not tune their background match to specific microhabitats, but instead, and regardless of treatment, all de…

Kuuleeko eduskunta tieteentekijöitä?

Viimeisen neljännesvuosisadan aikana eduskunnan valiokunnat kuulivat yli 10 000 kertaa tieteentekijöitä asiantuntijoina. Tutkijoiden osuus kaikista valiokuntien kuulemista henkilöstä on kuitenkin alle kymmenen prosenttia. Lisäksi tieteentekijöiden kuulemiset jakautuivat epätasaisesti valiokuntien, hallituskausien ja kuultujen tieteentekijöiden välillä. nonPeerReviewed

Variation in predator species abundance can cause variable selection pressure on warning signaling prey

Predation pressure is expected to drive visual warning signals to evolve toward conspicuousness. However, coloration of defended species varies tremendously and can at certain instances be considered as more camouflaged rather than conspicuous. Recent theoretical studies suggest that the variation in signal conspicuousness can be caused by variation (within or between species) in predators' willingness to attack defended prey or by the broadness of the predators' signal generalization. If some of the predator species are capable of coping with the secondary defenses of their prey, selection can favor reduced prey signal conspicuousness via reduced detectability or recognition. In this study…

Changes in predator community structure shifts the efficacy of two warning signals in Arctiid moths

Summary 1. Polymorphism in warning coloration is puzzling because positive frequency-dependent selection by predators is expected to promote monomorphic warning signals in defended prey. 2. We studied predation on the warning-coloured wood tiger moth (Parasemia plantaginis )b y using artificial prey resembling white and yellow male colour morphs in five separate populations with different naturally occurring morph frequencies. 3. We tested whether predation favours one of the colour morphs over the other and whether that is influenced either by local, natural colour morph frequencies or predator community composition. 4. We found that yellow specimens were attacked less than white ones rega…

Data from: Colour polymorphism torn apart by opposing positive frequency-dependent selection, yet maintained in space

1. Polymorphic warning signals in aposematic species are enigmatic because predator learning and discrimination should select for the most common coloration, resulting in positive frequency-dependent survival selection. 2. Here, we investigated whether differential mating success could create sufficiently strong negative frequency-dependent selection for rare morphs to explain polymorphic (white and yellow) warning coloration in male wood tiger moths (Parasemia plantaginis). 3. We conducted an experiment in semi-natural conditions where we estimated mating success for both white and yellow male moths under three different morph frequencies. 4. Contrary to expectations, mating success was po…

Data from: Temporal relationship between genetic and warning signal variation in the aposematic wood tiger moth (Parasemia plantaginis)

Many plants and animals advertise unpalatability through warning signals in the form of colour and shape. Variation in warning signals within local populations is not expected because they are subject to directional selection. However, mounting evidence of warning signal variation within local populations suggests that other selective forces may be acting. Moreover, different selective pressures may act on the individual components of a warning signal. At present, we have a limited understanding about how multiple selection processes operate simultaneously on warning signal components, and even less about their temporal and spatial dynamics. Here, we examined temporal variation of several w…

Data from: To quiver or to shiver: increased melanisation benefits thermoregulation, but reduces warning signal efficacy in the wood tiger moth

Melanin production is often considered costly, yet beneficial for thermoregulation. Studies of variation in melanization and the opposing selective forces that underlie its variability contribute greatly to understanding natural selection. We investigated whether melanization benefits are traded off with predation risk to promote observed local and geographical variation in the warning signal of adult male wood tiger moths (Parasemia plantaginis). Warning signal variation is predicted to reduce survival in aposematic species. However, in P. plantaginis, male hindwings are either yellow or white in Europe, and show continuous variation in melanized markings that cover 20 to 90 per cent of th…

Data from: Transparency reduces predator detection in mimetic clearwing butterflies

1. Predation is an important selective pressure and some prey have evolved conspicuous warning signals that advertise unpalatability (i.e. aposematism) as an antipredator defence. Conspicuous colour patterns have been shown effective as warning signals, by promoting predator learning and memory. Unexpectedly, some butterfly species from the unpalatable tribe Ithomiini possess transparent wings, a feature rare on land but common in water, known to reduce predator detection. 2. We tested if transparency of butterfly wings was associated with decreased detectability by predators, by comparing four butterfly species exhibiting different degrees of transparency, ranging from fully opaque to larg…

Supplementary data for article: Camouflage accuracy in Sahara-Sahel desert rodents

Antipredatory Function of Head Shape for Vipers and Their Mimics

Most research into the adaptive significance of warning signals has focused on the colouration and patterns of prey animals. However, behaviour, odour and body shape can also have signal functions and thereby reduce predators' willingness to attack defended prey. European vipers all have a distinctive triangular head shape; and they are all venomous. Several non-venomous snakes, including the subfamily Natricinae, commonly flatten their heads (also known as head triangulation) when disturbed. The adaptive significance of this potential behavioural mimicry has never been investigated. We experimentally tested if the triangular head shape typical of vipers offers protection against predation.…

Supplementary data for: Improved camouflage through ontogenetic colour change confers reduced detection risk in shore crabs

Many animals change appearance with age but the reasons why are rarely tested. Common shore crabs (Carcinus maenas), for example, are known for their ability to change colour over time. Young crabs show remarkable variation in coloration and it has been suggested that their variable appearance may help them to hide from predators in the habitats they use. However, as crabs grow they become more mobile and adult crabs, in contrast, are known to possess a more uniform coloration. This creates a problem: how to remain hidden in habitats that are variable and very different in appearance? To answer this, we first reared young shore crabs of two shades, pale or dark, on two background types rese…