Predator selection on multicomponent warning signals in an aposematic moth.

Aposematic prey advertise their unprofitability with conspicuous warning signals that are often composed of multiple color patterns. Many species show intraspecific variation in these patterns even though selection is expected to favor invariable warning signals that enhance predator learning. However, if predators acquire avoidance to specific signal components, this might relax selection on other aposematic traits and explain variability. Here, we investigated this idea in the aposematic moth Amata nigriceps that has conspicuous black and orange coloration. The size of the orange spots in the wings is highly variable between individuals, whereas the number and width of orange abdominal stripes remains consistent. We produced artificial moths that varied in the proportion of orange in the wings or the presence of abdominal stripes. We presented these to a natural avian predator, the noisy miner (Manorina melanocephala), and recorded how different warning signal components influenced their attack decisions. When moth models had orange stripes on the abdomen, birds did not discriminate between different wing signals. However, when the stripes on the abdomen were removed, birds chose the model with smaller wing spots. In addition, we found that birds were more likely to attack moths with a smaller number of abdominal stripes. Together, our results suggest that bird predators primarily pay attention to the abdominal stripes of A. nigriceps, and this could relax selection on wing coloration. Our study highlights the importance of considering individual warning signal components if we are to understand how predation shapes selection on prey warning coloration.

The protective value of the colour and shape of the mountain katydid's antipredator defence.

Deimatic behaviour is performed by prey when attacked by predators as part of an antipredator strategy. The behaviour is part of a sequence that consists of several defences, for example they can be preceded by camouflage and followed by a hidden putatively aposematic signal that is only revealed when the deimatic behaviour is performed. When displaying their hidden signal, mountain katydids (Acripeza reticulata) hold their wings vertically, exposing striking red and black stripes with blue spots and oozing an alkaloid-rich chemical defence derived from its Senecio diet. Understanding differences and interactions between deimatism and aposematism has proven problematic, so in this study we isolated the putative aposematic signal of the mountain katydid's antipredator strategy to measure its survival value in the absence of their deimatic behaviour. We manipulated two aspects of the mountain katydid's signal, colour pattern and whole body shape during display. We deployed five kinds of clay models, one negative control and four katydid-like treatments, in 15 grids across part of the mountain katydid's distribution to test the hypothesis that their hidden signal is aposematic. If this hypothesis holds true, we expected that the models, which most closely resembled real katydids would be attacked the least. Instead, we found that models that most closely resembled real katydids were the most likely to be attacked. We suggest several ideas to explain these results, including that the deimatic phase of the katydid's display, the change from a camouflaged state to exposing its hidden signal, may have important protective value.

A synthesis of deimatic behaviour.

Deimatic behaviours, also referred to as startle behaviours, are used against predators and rivals. Although many are spectacular, their proximate and ultimate causes remain unclear. In this review we aim to synthesise what is known about deimatic behaviour and identify knowledge gaps. We propose a working hypothesis for deimatic behaviour, and discuss the available evidence for the evolution, ontogeny, causation, and survival value of deimatic behaviour using Tinbergen's Four Questions as a framework. Our overarching aim is to direct future research by suggesting ways to address the most pressing questions in this field.

Recommendations for empowering early career researchers to improve research culture and practice.

Early career researchers (ECRs) are important stakeholders leading efforts to catalyze systemic change in research culture and practice. Here, we summarize the outputs from a virtual unconventional conference (unconference), which brought together 54 invited experts from 20 countries with extensive experience in ECR initiatives designed to improve the culture and practice of science. Together, we drafted 2 sets of recommendations for (1) ECRs directly involved in initiatives or activities to change research culture and practice; and (2) stakeholders who wish to support ECRs in these efforts. Importantly, these points apply to ECRs working to promote change on a systemic level, not only those improving aspects of their own work. In both sets of recommendations, we underline the importance of incentivizing and providing time and resources for systems-level science improvement activities, including ECRs in organizational decision-making processes, and working to dismantle structural barriers to participation for marginalized groups. We further highlight obstacles that ECRs face when working to promote reform, as well as proposed solutions and examples of current best practices. The abstract and recommendations for stakeholders are available in Dutch, German, Greek (abstract only), Italian, Japanese, Polish, Portuguese, Spanish, and Serbian.

Genetic and environmental drivers of colour and pattern in the Australian jacky dragon (Amphibolurus muricatus).

The underlying drivers of variation in the colouration (colour and pattern) of animals can be genetic, non-genetic, or more likely, a combination of both. Understanding the role of heritable genetic elements, as well as non-genetic factors such as age, habitat or temperature, in shaping colouration can provide insight into the evolution and function of these traits, as well as the speed of response to changing environments. This project examined the genetic and non-genetic drivers of continuous variation in colouration in a lizard, the jacky dragon (Amphibolurus muricatus). We leveraged a large captive experiment that manipulated parental and offspring thermal environment to simultaneously estimate the genetic and non-genetic drivers of variation in colouration. We found that the overall brightness, the elongation of the longitudinal stripes on the dorsum and the contrast between light and dark patches of the pattern were all heritable. Colouration varied according to the age of the hatchling; however, the thermal environment of neither the parents nor offspring contributed significantly to colouration. It appears that developmental plasticity and maternal effects associated with temperature are not important drivers of variation in our measures of colouration.

Additive genetic variation, but not temperature, influences warning signal expression in Amata nigriceps moths (Lepidoptera: Arctiinae).

Many aposematic species show variation in their color patterns even though selection by predators is expected to stabilize warning signals toward a common phenotype. Warning signal variability can be explained by trade-offs with other functions of coloration, such as thermoregulation, that may constrain warning signal expression by favoring darker individuals. Here, we investigated the effect of temperature on warning signal expression in aposematic Amata nigriceps moths that vary in their black and orange wing patterns. We sampled moths from two flight seasons that differed in the environmental temperatures and also reared different families under controlled conditions at three different temperatures. Against our prediction that lower developmental temperatures would reduce the warning signal size of the adult moths, we found no effect of temperature on warning signal expression in either wild or laboratory-reared moths. Instead, we found sex- and population-level differences in wing patterns. Our rearing experiment indicated that ~70% of the variability in the trait is genetic but understanding what signaling and non-signaling functions of wing coloration maintain the genetic variation requires further work. Our results emphasize the importance of considering both genetic and plastic components of warning signal expression when studying intraspecific variation in aposematic species.

The exploitation of sexual signals by predators: a meta-analysis.

Sexual signals are often central to reproduction, and their expression is thought to strike a balance between advertising to mates and avoiding detection by predatory eavesdroppers. Tests of the predicted predation costs have produced mixed results, however. Here we synthesized 187 effects from 78 experimental studies in a meta-analytic test of two questions; namely, whether predators, parasites and parasitoids express preferences for the sexual signals of prey, and whether sexual signals increase realized predation risk in the wild. We found that predators and parasitoids express strong and consistent preferences for signals in forced-choice contexts. We found a similarly strong overall increase in predation on sexual signallers in the wild, though here it was modality specific. Olfactory and acoustic signals increased the incidence of eavesdropping relative to visual signals, which experienced no greater risk than controls on average. Variation in outcome measures was universally high, suggesting that contexts in which sexual signalling may incur no cost, or even reduce the incidence of predation, are common. Our results reveal unexpected complexity in a central viability cost to sexual signalling, while also speaking to applied problems in invasion biology and pest management where signal exploitation holds promise for bio-inspired solutions.

Ecological responses to variation in seasonal snow cover.

Seasonal snow is among the most important factors governing the ecology of many terrestrial ecosystems, but rising global temperatures are changing snow regimes and driving widespread declines in the depth and duration of snow cover. Loss of the insulating snow layer will fundamentally change the environment. Understanding how individuals, populations, and communities respond to different snow conditions is thus essential for predicting and managing future ecosystem change. We synthesized 365 studies that examined ecological responses to variation in winter snow conditions. This research encompasses a broad range of methods (experimental manipulations, measurement of natural snow gradients, and long-term monitoring), locations (35 countries), study organisms (plants, mammals, arthropods, birds, fish, lichen, and fungi), and response measures. Earlier snowmelt was consistently associated with advanced spring phenology in plants, mammals, and arthropods. Reduced snow depth often increased mortality or physical injury in plants, although there were few clear effects on animals. Neither snow depth nor snowmelt timing had clear or consistent directional effects on body size of animals or biomass of plants. However, because 96% of studies were from the northern hemisphere, the generality of these trends across ecosystems and localities is also unclear. We identified substantial research gaps for several taxonomic groups and response types; research on wintertime responses was notably scarce. Future research should prioritize examination of the mechanisms underlying responses to changing snow conditions and the consequences of those responses for seasonally snow-covered ecosystems.

Respuestas Ecológicas a la Variación de la Cobertura Estacional de Nieve Resumen La nieve estacional se encuentra entre los factores más importantes que determinan la ecología de muchos ecosistemas terrestres, pero las crecientes temperaturas mundiales están cambiando los sistemas de nieve y causando declinaciones generalizadas en la profundidad y la duración de la capa de nieve. La pérdida de la capa de nieve aislante cambiará fundamentalmente el ambiente. El entendimiento de cómo los individuos, las poblaciones y las comunidades responden a las diferentes condiciones de nieve es esencial para predecir y manejar los cambios del ecosistema en el futuro. Sintetizamos 365 estudios que examinaron las respuestas ecológicas a la variación en las condiciones invernales de nieve. Esta investigación engloba una gama amplia de métodos (manipulaciones experimentales, medida de los gradientes naturales de nieve y monitoreo a largo plazo), localidades (35 países), organismos de estudio (plantas, mamíferos, artrópodos, aves, peces, líquenes y hongos) y medidas de respuesta. El deshielo temprano estuvo asociado continuamente con el adelanto de la fenología de plantas, mamíferos y artrópodos en primavera. La reducción de la profundidad de la nieve con frecuencia incrementó la mortalidad o las lesiones físicas en las plantas, aunque tuvo pocos efectos visibles sobre los animales. Ni la profundidad de la nieve ni la temporalidad del deshielo tuvieron efectos direccionales claros o consistentes sobre el tamaño corporal de los animales o la biomasa de las plantas. Sin embargo, ya que el 96% de los estudios se realizó en el hemisferio norte, la generalidad de estas tendencias en todos los ecosistemas y localidades tampoco está clara. Identificamos vacíos importantes en la investigación en torno a varios grupos taxonómicos y los tipos de respuesta; la información sobre las respuestas invernales estaba particularmente reducida. Las futuras investigaciones deberían priorizar el análisis de los mecanismos subyacentes a las respuestas ante las condiciones cambiantes de nieve y las consecuencias de aquellas respuestas para los ecosistemas cubiertos de nieve.

Taxonomic notes on the Australian skyhopper genus Kosciuscola Sjstedt (Orthoptera: Acrididae: Oxyinae).

The Australian skyhopper genus Kosciuscola Sjstedt consists of brachypterous species that inhabit the Australian alpine and subalpine region. The genus used to include 5 species and 1 subspecies, but according to a recent phylogenomic study, there could be as many as 14 species in the genus, that are genetically and geographically isolated from each other. This study represents the first step in describing and documenting the diversity of this interesting genus. In this study, we redefine the type species K. tristis, and elevate its subspecies K. tristis restrictus as a valid species on the basis of distinct morphological traits, geographical isolation, and phylogenomic evidence.

Predicting species and community responses to global change using structured expert judgement: An Australian mountain ecosystems case study.

Conservation managers are under increasing pressure to make decisions about the allocation of finite resources to protect biodiversity under a changing climate. However, the impacts of climate and global change drivers on species are outpacing our capacity to collect the empirical data necessary to inform these decisions. This is particularly the case in the Australian Alps which have already undergone recent changes in climate and experienced more frequent large-scale bushfires. In lieu of empirical data, we use a structured expert elicitation method (the IDEA protocol) to estimate the change in abundance and distribution of nine vegetation groups and 89 Australian alpine and subalpine species by the year 2050. Experts predicted that most alpine vegetation communities would decline in extent by 2050; only woodlands and heathlands are predicted to increase in extent. Predicted species-level responses for alpine plants and animals were highly variable and uncertain. In general, alpine plants spanned the range of possible responses, with some expected to increase, decrease or not change in cover. By contrast, almost all animal species are predicted to decline or not change in abundance or elevation range; more species with water-centric life-cycles are expected to decline in abundance than other species. While long-term ecological data will always be the gold standard for informing the future of biodiversity, the method and outcomes outlined here provide a pragmatic and coherent basis upon which to start informing conservation policy and management in the face of rapid change and a paucity of data.

Meta-analytic evidence for quantitative honesty in aposematic signals.

The combined use of noxious chemical defences and conspicuous warning colours is a ubiquitous anti-predator strategy. That such signals advertise the presence of defences is inherent to their function, but their predicted potential for quantitative honesty-the positive scaling of signal salience with the strength of protection-is the subject of enduring debate. Here, we systematically synthesized the available evidence to test this prediction using meta-analysis. We found evidence for a positive correlation between warning colour expression and the extent of chemical defences across taxa. Notably, this relationship held at all scales; among individuals, populations and species, though substantial between-study heterogeneity remains unexplained. Consideration of the design of signals revealed that all visual features, from colour to contrast, were equally informative of the extent of prey defence. Our results affirm a central prediction of honesty-based models of signal function and narrow the scope of possible mechanisms shaping the evolution of aposematism. They suggest diverse pathways to the encoding and exchange of information, while highlighting the need for deeper knowledge of the ecology of chemical defences to enrich our understanding of this widespread anti-predator adaptation.

The evolution of startle displays: a case study in praying mantises.

Anti-predator defences are typically regarded as relatively static signals that conceal prey or advertise their unprofitability. However, startle displays are complex performances that deter or confuse predators and can include a spectacular array of movements, colours and sounds. Yet, we do not fully understand the mechanisms by which they function, their evolutionary correlates, or the conditions under which they are performed and evolve. Here, we present, to our knowledge, the first phylogenetically controlled comparative analyses of startle displays including behavioural data, using praying mantises as a model system. We included 58 species that provide a good representation of mantis diversity and estimated the strength of phylogenetic signal in the presence and complexity of displays. We also tested hypotheses on potential evolutionary correlates, including primary defences and body size. We found that startle displays and morphological traits were phylogenetically conserved, whereas behavioural traits were highly labile. Surprisingly, body size was not correlated with display presence or complexity in phylogenetically controlled analyses. Species-rich clades were more likely to exhibit displays, suggesting that startle displays were probably involved in lineage diversification. We suggest that to further elucidate the conditions under which startle displays evolve, future work should include quantitative descriptions of multiple display components, habitat type, and predator communities. Understanding the evolution of startle displays is critical to our overall understanding of the theory behind predator-prey dynamics.

The protective value of a defensive display varies with the experience of wild predators.

Predation has driven the evolution of diverse adaptations for defence among prey, and one striking example is the deimatic display. While such displays can resemble, or indeed co-occur with, aposematic 'warning' signals, theory suggests deimatic displays may function independently of predator learning. The survival value of deimatic displays against wild predators has not been tested before. Here we used the mountain katydid Acripeza reticulata to test the efficacy of a putative deimatic display in the wild. Mountain katydids have a complex defence strategy; they are camouflaged at rest, but reveal a striking red-, blue-, and black-banded abdomen when attacked. We presented live katydids to sympatric (experienced) and allopatric (naive) natural predators, the Australian magpie Cracticus tibicen, and observed bird reactions and katydid behaviors and survival during repeated interactions. The efficacy of the katydids' defence differed with predator experience. Their survival was greatest when faced with naïve predators, which provided clear evidence of the protective value of the display. In contrast, katydid survival was consistently less likely when facing experienced predators. Our results suggest that sympatric predators have learned to attack and consume mountain katydids despite their complex defense, and that their post-attack display can be an effective deterrent, particularly against naïve predators. These results suggest that deimatism does not require predator learning to afford protection, but that a predator can learn to expect the display and subsequently avoid it or ignore it. That sympatric predators learn to ignore the defense is a possible explanation for the mountain katydid's counter-intuitive behavior of revealing warning colors only after tactile stimuli from predator attack.

The search for loci under selection: trends, biases and progress.

Detecting genetic variants under selection using FST outlier analysis (OA) and environmental association analyses (EAAs) are popular approaches that provide insight into the genetic basis of local adaptation. Despite the frequent use of OA and EAA approaches and their increasing attractiveness for detecting signatures of selection, their application to field-based empirical data have not been synthesized. Here, we review 66 empirical studies that use Single Nucleotide Polymorphisms (SNPs) in OA and EAA. We report trends and biases across biological systems, sequencing methods, approaches, parameters, environmental variables and their influence on detecting signatures of selection. We found striking variability in both the use and reporting of environmental data and statistical parameters. For example, linkage disequilibrium among SNPs and numbers of unique SNP associations identified with EAA were rarely reported. The proportion of putatively adaptive SNPs detected varied widely among studies, and decreased with the number of SNPs analysed. We found that genomic sampling effort had a greater impact than biological sampling effort on the proportion of identified SNPs under selection. OA identified a higher proportion of outliers when more individuals were sampled, but this was not the case for EAA. To facilitate repeatability, interpretation and synthesis of studies detecting selection, we recommend that future studies consistently report geographical coordinates, environmental data, model parameters, linkage disequilibrium, and measures of genetic structure. Identifying standards for how OA and EAA studies are designed and reported will aid future transparency and comparability of SNP-based selection studies and help to progress landscape and evolutionary genomics.

Deimatism: a neglected component of antipredator defence.

Deimatic or 'startle' displays cause a receiver to recoil reflexively in response to a sudden change in sensory input. Deimatism is sometimes implicitly treated as a form of aposematism (unprofitability associated with a signal). However, the fundamental difference is, in order to provide protection, deimatism does not require a predator to have any learned or innate aversion. Instead, deimatism can confer a survival advantage by exploiting existing neural mechanisms in a way that releases a reflexive response in the predator. We discuss the differences among deimatism, aposematism, and forms of mimicry, and their ecological and evolutionary implications. We highlight outstanding questions critical to progress in understanding deimatism.

Within-season variability of fighting behaviour in an Australian alpine grasshopper.

Throughout the breeding season, changing environmental and biological conditions can lead to variation in the reproductive landscape of many species. In alpine environments temperature is a key driver of behaviour for small ectotherms such as insects, but variable biotic factors such as mate quality and availability can also influence behaviour. Kosicuscola tristis is a small semelparous grasshopper of the Australian alpine region. In a rare behaviour among grasshoppers, K. tristis males engage in vigorous fights over access to females, involving mandible displays, kicking, biting and grappling. In this study we describe the variation in fighting behaviour of K. tristis throughout the breeding season and test several hypotheses related to temperature, body size, mating behaviour, and female quality. We show that K. tristis males are more aggressive toward each other at the end of the breeding season than at the beginning. This increased aggression is associated with decreased daily average temperatures (from ~20°C to ~9°C), decreased mating activity, increased female fecundity, and an unexpected trend toward an increase in female-to-male aggression. These results suggest that K. tristis is likely under increased selective pressure to time key life cycle events with favourable biological and climatic conditions. The stochastic nature of alpine environments combined with a relatively short life span and breeding season, as well as limited mating opportunities toward the end of the season may have contributed to the evolution of this extraordinary mating system.

Bioluminescence.

Kahle and Umbers introduce the ways by which organisms emit light though chemical reactions.

The mothematics of female pheromone signaling: strategies for aging virgins.

Although females rarely experience strong mate limitation, delays or lifelong problems of mate acquisition are detrimental to female fitness. In systems where males search for females via pheromone plumes, it is often difficult to assess whether female signaling is costly. Direct costs include the energetics of pheromone production and attention from unwanted eavesdroppers, such as parasites, parasitoids, and predators. Suboptimal outcomes are also possible from too many or too few mating events or near-simultaneous arrival of males who make unwanted mating attempts (even if successfully thwarted). We show that, in theory, even small costs can lead to a scenario where young females signal less intensely (lower pheromone concentration and/or shorter time spent signaling) and increase signaling effort only as they age and gather evidence (while still virgin) on whether sperm limitation threatens their reproductive success. Our synthesis of the empirical data available on Lepidoptera supports this prediction for one frequently reported component of signaling-time spent calling (often reported as the time of onset of calling at night)-but not for another, pheromone titer. This difference is explicable under the plausible but currently untested assumption that signaling earlier than other females each night is a more reliable way of increasing the probability of acquiring at least one mate than producing a more concentrated pheromone plume.