Biology subjects

Nokelainen, O.

Publications and source records attributed to Nokelainen, O..

8 recordsLinked to original sources

bioRxiv2026

Resolution of multi-receiver trade-offs in wall lizard colouration

Animal colouration evolves under multiple, often conflicting, selective pressures. Conspicuous, non-aposematic colour-patterns that enhance conspecific communication may simultaneously increase detectability by predators. Such trade-offs can be resolved by optimising colour-patterns to match the perceptual abilities of different receivers. We tested whether dorsal colour-patterns of the Lusitanian wall lizard (Podarcis lusitanicus) are optimised for ecologically relevant receivers across relevant viewing distances, while accounting for the visual acuity of conspecifics and predators. Conspecifics and snakes detected chromatic information at shorter distances, whereas achromatic and luminance information were detected at longer distances. Birds showed a uniform decline in detectability across the colour-pattern components with increasing viewing distance. Larger males retained high chromatic detectability across all receivers despite the general distance-related decline, whereas females and smaller individuals exhibited less salient colour patterns, consistent with predator avoidance strategy. Our results show that lizards resolve the trade-off between conspecific communication and predator detection through distance-dependent colour-pattern perceptibility across receivers. This resolution breaks down in large males, for whom the benefits of salient chromatic patterns for intraspecific communication may outweigh increased detectability to predators.

evolutionary biology↗

bioRxiv2026

Independent major-effect loci and limited trait associations characterise warning colour variation across sexes and life stages in the wood tiger moth (Arctia plantaginis)

Variation in warning colouration is common, despite strong positive frequency-dependent selection favouring the most common morph, and may arise from spatially or temporally variable selection or associations with other traits. We investigated the maintenance of warning colour variation in the wood tiger moth, Arctia plantaginis, which shows sexual dimorphism in colouration: two distinct male morphs controlled by a single locus, and continuous variation in female hindwing colour and larval warning signal size. Using whole-genome sequencing of 657 individuals from a laboratory stock derived from a wild polymorphic population, we examined the genetic basis of multiple life-history traits, chemical defence, pheromones, and male activity. We specifically tested for associations with male colour and aimed to identify the genetic basis of female and larval colour variation. We detected separate major-effect loci for female colour and larval signal size, which were different from the male colour locus. Hindwing melanism was the only trait associated with male colour, mapping to a nearby major-effect locus and showing a link to reduced activity at higher temperatures. Female colour was phenotypically correlated with pheromone amount. However, apart from melanism, none of the other traits measured showed an association with any of the loci controlling warning colours and we found no evidence that male colour is part of a supergene or complex co-adapted phenotype. Instead, our results indicate largely independent genetic architectures for warning colours suggesting that variable selection rather than genetic associations may contribute to the maintenance of colour polymorphisms in this species.

evolutionary biology↗

bioRxiv2025

Biologically inspired warning patterns deter birds from wind turbines

Wind power has been at the forefront of investment and innovation in renewable energy. However, bird fatalities from collisions with wind turbines present an ecological and social challenge to the growing deployment of wind power. Increasingly, research has focused on utilising the sensory ecology of animals to provide passive or active cues that minimise collision risk by increasing the detectability and/or aversiveness of the turbine blades and towers. In nature, numerous aposematic species use contrasting colours and striped patterns to warn birds of their unprofitability. These common signal elements are effective due to their salience within a wide range of natural scenes, memorability, generalisability across taxa due to mimicry, and exploitation of the innate colour preferences of birds. This begs the question: might biologically inspired turbine blades that mimic aposematic patterns help protect birds by increasing their avoidance of turbine blades? Here, we used a screen-based game experimental setup to test the behavioural responses of wild-caught great tits (Parus major) to three existing wind turbine patterns (white, red striped, and single black blade) as well as a novel biologically inspired aposematic pattern. Birds were less likely to approach and, when they did approach, took significantly longer to approach patterned compared to uniform white blades. This effect was strongest for our aposematic pattern compared to all other patterns tested, highlighting the utility of our bio-inspired approach. Our work suggests that adding red, black and yellow warning patterns to wind turbine blades could reduce bird collisions with wind turbines.

ecology↗

bioRxiv2024

Honest signalling in predator-prey interactions: testing the resource allocation hypothesis

Warning signals are honest if they reliably deliver information about prey unprofitability to predators. One potential mechanism that may create and maintain a positive relationship between the strength of signals and defence is the resource allocation between these costly traits. Here, we test this hypothesis using the wood tiger moth Arctia plantaginis, whose females red hindwings are a warning signal to predators but show considerable variation in colouration within populations. These moths also produce a defensive chemical that is known to influence avian predator attack risk. Using dietary manipulations, image and chemical analyses, and experiments with ecologically relevant predators we demonstrate that protein availability during development can influence the strength of both the primary warning signal and the secondary defence. Our results show that females raised on a high-protein or ad libitum natural diet produced more distasteful defensive fluids than those raised on a low-protein diet or subjected to periodic food deprivation. While the patterning of the warning signal was unaffected by food deprivation, its efficacy was diminished in moths raised on a low-protein diet. However, this change was imperceptible to avian predators. Critically, resource availability influenced the relationship between signal strength and defence: moths on a high-protein diet displayed a positive correlation between warning signal strength and unpalatability, whereas this correlation was absent in moths raised on a natural diet. These findings show that resource availability can weaken the reliability of warning signals as an indicator of an individuals defensive capabilities, highlighting the complex interplay between ecological conditions and the evolution of honest signalling.

evolutionary biology↗

bioRxiv2023

Positive and negative frequency-dependent selection acting on polymorphism in a palatable moth

Camouflage and warning signals are contrasted prey strategies reducing predator attack, which offer an excellent opportunity to study the evolutionary forces acting on prey appearance. Edible prey are often inconspicuous and escape predation by remaining undetected. Predators learn to find the most common ones, leading to apostatic selection (advantage to rare morphs) enhancing variation in cryptic prey. By contrast, defended prey are often conspicuous and escape predation by using warning colorations identifying them as unprofitable. Predators avoid the ones they are most familiar with, leading to positive frequency-dependence and warning signal uniformity. It is less clear, however, what happens when two morphs of the same species vary strongly in conspicuousness, and how to explain the maintenance of cryptic and conspicuous morphs within populations, in the case of profitable prey. Using the white and melanic morphs of the invasive Box Tree Moth (Cydalima perspectalis) presented at three different frequencies, we investigate whether a) caterpillars and adult moths are palatable for birds, b) the less conspicuous, melanic morph experiences lower predation rates and b) whether frequency-dependence balances morph frequencies. Our results suggest that the melanic morph enjoys a survival advantage owing to a lower visibility. However, our experiments show that, unexpectedly, the two color morphs experience opposite patterns of frequency-dependent predation, despite being both fully palatable to birds. The melanic morph is under apostatic selection, whereas the conspicuous, white morph is subject to positive frequency-dependence (safety in numbers). Our experiments also show some level of unpalatability in the caterpillars. These results offer novel insight into how predation triggers contrasting evolutionary patterns in a palatable, polymorphic species within two morphs that differ markedly in conspicuousness and within two different life stages. Lay summaryUnderstanding the factors influencing character variation in natural populations is a key question in evolutionary ecology. Predation is one of the main drivers of color evolution in prey communities and prey usually mitigate predation using camouflage or warning colors. Camouflage evolves because it lowers the probability of being detected by predators. Since predators are more efficient at finding prey which they are familiar with, prey which display a rare phenotype are favoured (negative frequency-dependent selection). By contrast, aposematism is defined by conspicuous appearance in toxic or otherwise unprofitable prey, and evolves because birds identify defended prey by learning to use their appearance as a warning signal. The most common signals are usually best identified and avoided (positive-frequency dependent selection). It is not clear, however, how these two forces combine when predators are facing cryptic and conspicuous morphs of the same species, and how to explain their coexistence. Here we investigate this question in a laboratory experiment, by presenting wild birds with a melanic and a white morph of the same moth. Unexpectedly, our results show that despite being both fully palatable to birds, the two color morphs are subject to very different types of selection depending on their frequencies. The melanic morph is favored when it is rare, the conspicuous white morph as it gets common. The simultaneous action of these forces may contribute to maintain color polymorphism in natural populations. We also show that caterpillars of this species are unpalatable and chemically defended, whereas adults are not, showing opposite strategies of predator defense in different life stages of the same species.

evolutionary biology↗

bioRxiv2023

Diet influences resource allocation in chemical defence but not melanin synthesis in an aposematic moth

For animals that synthesise their chemical compounds de novo, resources, particularly proteins, can influence investment in chemical defences and nitrogen-based wing colouration such as melanin. Competing for the same resources often leads to trade-offs in resource allocation. We manipulated protein availability in the larval diet of the wood tiger moth, Arctia plantaginis, to test how early life resource availability influences relevant life history traits, melanin production, and chemical defences. We expected higher dietary protein to result in more effective chemical defences and a higher amount of melanin in the wings. According to the resource allocation hypothesis, we also expected individuals with less melanin to have more resources to allocate to chemical defences. We found that protein-deprived moths had a slower larval development, and their chemical defences were less unpalatable for bird predators, but the expression of melanin in their wings did not differ from that of moths raised on a high-protein diet. The amount of melanin in the wings, however, unexpectedly correlated positively with chemical defences, irrespective of the diet they were raised on. Our findings demonstrate that the resources available in early life have an important role in the efficacy of chemical defences, but melanin-based warning colours are less sensitive to resource variability than other fitness-related traits.

ecology↗

bioRxiv2020

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

Warning 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 Georgia, where the predator community was the least diverse, whereas in the most diverse avian community in Estonia, hardly any models were attacked. Our results support the idea that spatial variation in predator and prey communities alters the strength or direction of selection on warning signals, thus facilitating a geographic mosaic of selection.

ecology↗

bioRxiv2018

Transparency reduces predator detection in chemically protected clearwing butterflies

1. 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.\n\n2. 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 experiments. Vision modelling predicted detectability to be similar for these two predator types.\n\n3. In concordance with predictions, more transparent species were almost never the first detected items and were detected less often than the opaque species by both birds and humans, suggesting that transparency enhances crypsis. However, humans could learn to better detect the most transparent species over time. Our study demonstrates for the first time that transparency on land likely decreases detectability by visual predators.

evolutionary biology↗