Search

The colours of insects function in intraspecific communication such as sexual signalling, interspecific communication such as protection from predators, and in physiological processes, such as thermoregulation. The expression of melanin-based colours is temperature-dependent and thus likely to be impacted by a changing climate. However, it is unclear how climate change drives changes in body and wing colour may impact insect physiology and their interactions with conspecifics (e.g. mates) or heterospecific (e.g. predators or prey). The aim of this review is to synthesise the current knowledge of the consequences of climate-driven colour change on insects. Here, we discuss the environmental factors that affect insect colours, and then we outline the adaptive mechanisms in terms of phenotypic plasticity and microevolutionary response. Throughout we discuss the impact of climate-related colour change on insect physiology, and interactions with con-and-heterospecifics., Competing Interests: The authors declare no competing interests., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

Adaptive evolution relies on both heritable variation and selection. Variation is the raw material upon which selection acts, so any mechanism that limits or prevents the generation of heritable variation reduces the power of selection to lead to adaptation. Such limitations are termed evolutionary constraints. While it is widely accepted that constraints play an important role in shaping evolutionary outcomes, their relative importance, as opposed to adaptation, in determining evolutionary outcomes remains a subject of debate. Evolutionary constraints are often evoked as the reason behind the persistence of inaccurate mimicry. Here, we compared the variation and accuracy of body-shape mimicry in ant-mimicking spiders with that of ant-mimicking insects, predicting greater constraints, and hence inaccuracy, in spiders mimicking ants, due to their evolutionary distance from the ant model. We found high inter-species variation in mimetic accuracy, but dorsally, no overall difference in mimetic accuracy between spider and insect mimics, which is inconsistent with a constraint causing inaccurate mimicry. Our study provides empirical evidence suggesting that imperfect mimicry in spiders and insects is predominantly shaped by adaptive processes rather than constraints or chance. Our findings contribute to our understanding of the mechanisms underlying evolutionary diversity and the processes that shape phenotypic outcomes.

Host-parasite interactions are impacted by climate, which may result in variation of parasitism across landscapes and time. Understanding how parasitism varies across these spatio-temporal scales is crucial to predicting how organisms will respond to and cope under a rapidly changing climate. Empirical work on how parasitism varies across climates is limited. Here, we examine the variation of parasitism across seasons and identify the likely climatic factors that explain this variation using Agriocnemis femina damselflies and Arrenurus water mite ectoparasites as a host-parasite study system. We assessed parasitism in a natural population in Sylhet, Bangladesh which is located in subtropical climate between 2021 and 2023. We calculated prevalence (proportion of infected individuals) and intensity (the number of parasites on an infected individual) of parasitism across different seasons. Parasite prevalence and intensity were greater during cooler seasons (autumn and winter) compared to hotter seasons (spring and summer). Mean temperature and precipitation were negatively correlated with parasite prevalence, whereas only mean precipitation was negatively correlated with parasite intensity. Tropical, subtropical and mediterranean regions are predicted to experience extreme climatic events (extreme temperature, less precipitation and frequent drought) as a consequence of anthropogenic climate change, and our finding suggests that this might alter patterns of parasitism in aquatic insects., Competing Interests: The authors declare no conflict of interest., (© 2024 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The current consensus is that sexual selection is responsible for the rapid and diverse evolution of genitalia, with several mutually exclusive mechanisms under debate, including non-antagonistic, antagonistic and stabilizing mechanisms. We used the orb-web spider, Argiope lobata (Araneidae), as a study model to quantify the allometric relationship between body size and genitalia, and to test for any impact of genital structures on male mating success or outcome in terms of copulation duration, leg loss or cannibalism. Our data do not support the 'one-size-fits-all' hypothesis that predicts a negative allometric slope between genitalia and body size. Importantly, we measured both male and female genitalia, and there was no sex specific pattern in allometric slopes. Unexpectedly, we found no predictor for reproductive success as indicated by copulation duration, cannibalism, and leg loss., Competing Interests: The authors declare there are no competing interests., (©2023 Dharmarathne et al.)

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 Author(s) 2023. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology.)

Prey seldom rely on a single type of antipredator defence, often using multiple defences to avoid predation. In many cases, selection in different contexts may favour the evolution of multiple defences in a prey. However, a prey may use multiple defences to protect itself during a single predator encounter. Such "defence portfolios" that defend prey against a single instance of predation are distributed across and within successive stages of the predation sequence (encounter, detection, identification, approach (attack), subjugation and consumption). We contend that at present, our understanding of defence portfolio evolution is incomplete, and seen from the fragmentary perspective of specific sensory systems (e.g., visual) or specific types of defences (especially aposematism). In this review, we aim to build a comprehensive framework for conceptualizing the evolution of multiple prey defences, beginning with hypotheses for the evolution of multiple defences in general, and defence portfolios in particular. We then examine idealized models of resource trade-offs and functional interactions between traits, along with evidence supporting them. We find that defence portfolios are constrained by resource allocation to other aspects of life history, as well as functional incompatibilities between different defences. We also find that selection is likely to favour combinations of defences that have synergistic effects on predator behaviour and prey survival. Next, we examine specific aspects of prey ecology, genetics and development, and predator cognition that modify the predictions of current hypotheses or introduce competing hypotheses. We outline schema for gathering data on the distribution of prey defences across species and geography, determining how multiple defences are produced, and testing the proximate mechanisms by which multiple prey defences impact predator behaviour. Adopting these approaches will strengthen our understanding of multiple defensive strategies., (© 2023 The Authors. Journal of Evolutionary Biology published by John Wiley & Sons Ltd on behalf of European Society for Evolutionary Biology.)

Tool-use in animals is a complex and rare phenomenon, particularly in insects. Tool-use in assassin bugs has been suggested as several species apply adhesive plant resins to their body, which has been hypothesized to function in enhancing prey capture. Here, we staged predatory interactions of resin-deprived and resin-equipped assassin bugs ( Gorareduvius sp.) and discovered that applying resin as a tool conveys a clear predatory advantage to the assassin bugs. Gorareduvius sp. can thus be considered a tool-user, and since this behaviour was present in all individuals, including newly hatched nymphs, tool-use can be considered to be stereotyped. Our study, along with others, suggests that, when compared with other insects, tool-use is disproportionately common within the assassin bugs.

Cooperation should only evolve if the direct and/or indirect benefits exceed the costs. Hence, cooperators are expected to generate selective benefits for themselves and the kin-group while defectors will impose costs. The subsocial spider, Australomisidia ergandros, shows consistent cooperation and defection tactics while foraging. Cooperative individuals are consistently likely to share prey with other group members whereas defector spiders rarely share the prey they acquired. Here, we assess costs and benefits of cooperation, and the causal determinants behind cooperative and defective phenotypes. We constructed experimental kin-colonies of A. ergandros composed of pure cooperative or defector foragers and show that pure cooperative groups had higher hunting success as they acquired prey more quickly with greater joint participation than pure defector groups. Importantly, defectors suffered higher mortality than cooperators and lost considerable weight. A social network approach using subadult spiders revealed that foraging tactic is sex dependent with males cooperating more frequently than females. Our results provide a rare empirical demonstration of sex-specific male cooperation that confer individual and kin-group benefits., (© 2023. The Author(s).)

A prominent question in animal research is how the evolution of morphology and ecology interacts in the generation of phenotypic diversity. Spiders are some of the most abundant arthropod predators in terrestrial ecosystems and exhibit a diversity of foraging styles. It remains unclear how spider body size and proportions relate to foraging style, and if the use of webs as prey capture devices correlates with changes in body characteristics. Here, we present the most extensive data set to date of morphometric and ecological traits in spiders. We used this data set to estimate the change in spider body sizes and shapes over deep time and to test if and how spider phenotypes are correlated with their behavioral ecology. We found that phylogenetic variation of most traits best fitted an Ornstein-Uhlenbeck model, which is a model of stabilizing selection. A prominent exception was body length, whose evolutionary dynamics were best explained with a Brownian Motion (free trait diffusion) model. This was most expressed in the araneoid clade (ecribellate orb-weaving spiders and allies) that showed bimodal trends toward either miniaturization or gigantism. Only few traits differed significantly between ecological guilds, most prominently leg length and thickness, and although a multivariate framework found general differences in traits among ecological guilds, it was not possible to unequivocally associate a set of morphometric traits with the relative ecological mode. Long, thin legs have often evolved with aerial webs and a hanging (suspended) locomotion style, but this trend is not general. Eye size and fang length did not differ between ecological guilds, rejecting the hypothesis that webs reduce the need for visual cue recognition and prey immobilization. For the inference of the ecology of species with unknown behaviors, we propose not to use morphometric traits, but rather consult (micro-)morphological characters, such as the presence of certain podal structures. These results suggest that, in contrast to insects, the evolution of body proportions in spiders is unusually stabilized and ecological adaptations are dominantly realized by behavioral traits and extended phenotypes in this group of predators. This work demonstrates the power of combining recent advances in phylogenomics with trait-based approaches to better understand global functional diversity patterns through space and time. [Animal architecture; Arachnida; Araneae; extended phenotype; functional traits; macroevolution; stabilizing selection.]., (© The Author(s) 2022. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Spiders, the most specious taxon of predators, have evolved an astounding range of predatory strategies, including group hunting, specialized silk traps, pheromone-loaded bolas, and aggressive mimicry. Spiders that hunt prey defended with behavioral, mechanical, or chemical means are under additional selection pressure to avoid injury and death. Ants are considered dangerous because they can harm or kill their predators, but some groups of spiders, such as the Theridiidae, have a very high diversification of ant-hunting species and strategies [J. Liu et al., Mol. Phylogenet. Evol . 94, 658-675 (2016)]. Here, we provide detailed behavioral analyses of the highly acrobatic Australian ant-slayer spider, Euryopis umbilicata (Theridiidae), that captures much larger and defended Camponotus ants on vertical tree trunks. The hunting sequence consists of ritualized steps performed within split seconds, resulting in an exceptionally high prey capture success rate.

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., Competing Interests: The authors have no conflicts of interest to declare., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The prevalence and intensity of parasitism can have different fitness costs between sexes, and across species and developmental stages. This variation could arise because of species specific sexual and developmental differences in body condition, immunity, and resistance. Theory predicts that the prevalence of parasitism will be greater in individuals with poor body condition and the intensity of parasitism will be greater in individuals with larger body size. These predictions have been tested and verified in vertebrates. In insects, however, contradictory evidence has been found in different taxa. Here, we tested these predictions on two species of Agriocnemis (Agriocnemis femina and Agriocnemis pygmaea) damselflies, which are parasitized by Arrenurus water mite ectoparasites. We measured body weight, total body length, abdomen area and thorax area of non-parasitized damselflies and found body condition varied between males and females, between immature females and mature females and between A. femina and A. pygmaea. Then, we calculated the parasite prevalence, i.e., the absence or presence of parasites and intensity, i.e., the number of parasites per infected damselfly in eleven natural populations of both species. In line to our predictions, we observed greater prevalence in immature females than mature females but found no difference in parasite prevalence between males and females. Furthermore, we found that parasite intensity was higher in females than males and in immature females than mature females. Our result also showed that the frequency and intensity of parasitism varied between the two studied species, being higher in A. pygmaea than A. femina. Our study provides evidence that parasitism impacts sexes, developmental stages and species differentially and suggests that variation may occur due to sex, developmental stage, and species-specific resistance and tolerance mechanism., Competing Interests: The authors have declared that no competing interests exist.

Trait databases have become important resources for large-scale comparative studies in ecology and evolution. Here we introduce the AnimalTraits database, a curated database of body mass, metabolic rate and brain size, in standardised units, for terrestrial animals. The database has broad taxonomic breadth, including tetrapods, arthropods, molluscs and annelids from almost 2000 species and 1000 genera. All data recorded in the database are sourced from their original empirical publication, and the original metrics and measurements are included with each record. This allows for subsequent data transformations as required. We have included rich metadata to allow users to filter the dataset. The additional R scripts we provide will assist researchers with aggregating standardised observations into species-level trait values. Our goals are to provide this resource without restrictions, to keep the AnimalTraits database current, and to grow the number of relevant traits in the future., (© 2022. The Author(s).)

Many ecological interactions of spiders with their potential prey and predators are affected by the visibility of their bodies and silk, especially in habitats with lower structural complexity that expose spiders. For instance, the surface of tree trunks harbours relatively limited structures to hide in and may expose residents to visual detection by prey and predators. Here we provide the first detailed description of the novel retreat building strategy of the tree trunk jumping spider Arasia mullion . Using fields surveys, we monitored and measured over 115 spiders and 554 silk retreats. These spiders build silk retreats on the exposed surface of tree trunks, where they remain as sedentary permanent residents. Furthermore, the spiders decorate the silk retreats with bark debris that they collect from the immediate surrounding. We discuss the role of silk decoration in the unusual sedentary behaviour of these spiders and the potential mechanisms that allow A. mullion to engineer their niche in a challenging habitat., Competing Interests: The authors declare that they have no competing interests., (© 2022 Aceves-Aparicio et al.)

AbstractUnderstanding Batesian mimicry is a classic problem in evolutionary biology. In Batesian mimicry, a defended species (the model) is mimicked by an undefended species (the mimic). Prior theories have emphasized the role of predator behavior and learning as well as evolution in model-mimic complexes but have not examined the role of population dynamics in potentially governing the relative abundances and even persistence of model-mimic systems. Here, we examined the effect of the population dynamics of predators and alternative prey on the prevalence of warning-signaling prey composed of models and mimics. Using optimal foraging theory and signal detection theory, we found that the inclusion of predator and alternative prey population dynamics could reverse traditional theoretical predictions: as alternative prey increase in numbers, mimics suffer because larger populations of predators are maintained, resulting in apparent competition. Under some circumstances, apparent competition affects model populations as well, although not as severely as it affects mimics. Our results bear on the intriguing puzzle that in nature warning signals are relatively scarce, yet experiments suggest that such signals can be highly advantageous. The availability of alternative prey and numerical responses by predators can overwhelm advantages observed in experiments to keep warning signals in model-mimic systems relatively scarce.

Warning signals are a striking example of natural selection present in almost every ecological community - from Nordic meadows to tropical rainforests, defended prey species and their mimics ward off potential predators before they attack. Yet despite the wide distribution of warning signals, they are relatively scarce as a proportion of the total prey available, and more so in some biomes than others. Classically, warning signals are thought to be governed by positive density-dependent selection, i.e. they succeed better when they are more common. Therefore, after surmounting this initial barrier to their evolution, it is puzzling that they remain uncommon on the scale of the community. Here, we explore factors likely to determine the prevalence of warning signals in prey assemblages. These factors include the nature of prey defences and any constraints upon them, the behavioural interactions of predators with different prey defences, the numerical responses of predators governed by movement and reproduction, the diversity and abundance of undefended alternative prey and Batesian mimics in the community, and variability in other ecological circumstances. We also discuss the macroevolution of warning signals. Our review finds that we have a basic understanding of how many species in some taxonomic groups have warning signals, but very little information on the interrelationships among population abundances across prey communities, the diversity of signal phenotypes, and prey defences. We also have detailed knowledge of how a few generalist predator species forage in artificial laboratory environments, but we know much less about how predators forage in complex natural communities with variable prey defences. We describe how empirical work to address each of these knowledge gaps can test specific hypotheses for why warning signals exhibit their particular patterns of distribution. This will help us to understand how behavioural interactions shape ecological communities., (© 2021 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Spiders are a highly diversified group of arthropods and play an important role in terrestrial ecosystems as ubiquitous predators, which makes them a suitable group to test a variety of eco-evolutionary hypotheses. For this purpose, knowledge of a diverse range of species traits is required. Until now, data on spider traits have been scattered across thousands of publications produced for over two centuries and written in diverse languages. To facilitate access to such data, we developed an online database for archiving and accessing spider traits at a global scale. The database has been designed to accommodate a great variety of traits (e.g. ecological, behavioural and morphological) measured at individual, species or higher taxonomic levels. Records are accompanied by extensive metadata (e.g. location and method). The database is curated by an expert team, regularly updated and open to any user. A future goal of the growing database is to include all published and unpublished data on spider traits provided by experts worldwide and to facilitate broad cross-taxon assays in functional ecology and comparative biology. Database URL:https://spidertraits.sci.muni.cz/., (© The Author(s) 2021. Published by Oxford University Press.)

Do animals set the course for the evolution of their lineage when manipulating their environment? This heavily disputed question is empirically unexplored but critical to interpret phenotypic diversity. Here, we tested whether the macroevolutionary rates of body morphology correlate with the use of built artifacts in a megadiverse clade comprising builders and nonbuilders-spiders. By separating the inferred building-dependent rates from background effects, we found that variation in the evolution of morphology is poorly explained by artifact use. Thus natural selection acting directly on body morphology rather than indirectly via construction behavior is the dominant driver of phenotypic diversity., Competing Interests: The authors declare no competing interest., (Copyright © 2021 the Author(s). Published by PNAS.)

Many animals mimic dangerous or undesirable prey as a defence from predators. We would expect predators to reliably avoid animals that closely resemble dangerous prey, yet imperfect mimics are common across a wide taxonomic range. There have been many hypotheses suggested to explain imperfect mimicry, but comparative tests across multiple mimicry systems are needed to determine which are applicable, and which-if any-represent general principles governing imperfect mimicry. We tested four hypotheses on Australian ant mimics and found support for only one of them: the information limitation hypothesis. A predator with incomplete information will be unable to discriminate some poor mimics from their models. We further present a simple model to show that predators are likely to operate with incomplete information because they forage and make decisions while they are learning, so might never learn to properly discriminate poor mimics from their models. We found no evidence that one accurate mimetic trait can compensate for, or constrain, another, or that rapid movement reduces selection pressure for good mimicry. We argue that information limitation may be a general principle behind imperfect mimicry of complex traits, while interactions between components of mimicry are unlikely to provide a general explanation for imperfect mimicry.

Aggressive and cannibalistic female spiders can impose strong selection on male mating and fertilization strategies. Furthermore, the distinctive reproductive morphology of spiders is predicted to influence the outcome of sperm competition. Polyandry is common in spiders, leading to defensive male strategies that include guarding, plugging and self-sacrifice. Paternity patterns are highly variable and unlikely to be determined solely by mating order, but rather by relative copulation duration, deployment of plugs and cryptic female choice. The ability to strategically allocate sperm is limited, either by the need to refill pedipalps periodically or owing to permanent sperm depletion after mating. Further insights now rely on unravelling several proximate mechanisms such as the process of sperm activation and the role of seminal fluids. This article is part of the theme issue 'Fifty years of sperm competition'.

It has been suggested that physical interactions between biological and environmental surfaces may constrain ecological niche spaces. However, the mechanistic understanding of niche formation is frequently limited by the lack of information on the function and variation of these interactions. Here, we hypothesised that two closely related species of orb-web spiders have evolved different adhesion performance of web attachment (i.e. piriform silk) facilitating the occupation of contrasting microhabitats: plants versus rocks. Contrary to our prediction, we found that piriform silk adhesion was equally affected by surface chemistry in both species, with maximal adhesion on surfaces with high surface polarity and an average adhesion loss of 70-75% on low polar surfaces. Spiders did not respond to adhesion losses by increasing the anchor size, despite the repeated failure to attach their web to low polar surfaces. In a natural setting, poor adhesion on low polar surfaces may be mitigated by behavioural means, like the preference to place anchors on corrugated surface features such as leaf edges, or the spinning of multiple anchorages and formation of a bundled anchor line. Thus, microhabitat choice for web-building spiders may be governed by structural properties rather than surface chemistry. These results suggest that the repeatedly demonstrated effects of surface chemistry on bio-adhesion may be ecologically less important than assumed and that the role of behaviour in the evolution of bio-adhesion performance has been underestimated.

Background: Queensland fruit fly (Q-fly) is a destructive insect pest that infests a wide variety of agricultural plants in Australia. The sterile insect technique (SIT) is used to manage Q-flies, but the effectiveness of SIT has not been tested in the presence of natural predators. The objective of this study was to investigate the effect of natural predators and SIT on the survival and reproduction of laboratory reared Q-flies under semi-natural conditions. We altered the presence of predators and irradiated Q-fly males, and measured survival, number of eggs laid and egg-hatching rate., Results: The presence of natural predators significantly affected the survival of Q-flies and appeared to decrease the number of eggs laid. Interestingly, we found that both sterile and fertile males were more prone to predation than females, but we found no difference among males. The presence of sterile males significantly reduced Q-fly fertility, but the interaction of natural predators and sterile males did not significantly reduce the number of fertile eggs., Conclusion: Our findings highlight the important role of natural predators in controlling Q-flies together with SIT and provide a solid foundation for similar large-scale field trials using wild counterparts. © 2019 Society of Chemical Industry., (© 2019 Society of Chemical Industry.)

Animals are able to assess the risk of predation and respond accordingly via behavioural and physiological changes. Web-building spiders are in the unique situation where they reside in the middle of their web and are therefore relatively exposed to predators. Thus, these spiders might moderate either their web-building behaviour or their behaviour on the web when exposed to the threat of predation. In this study, we experimentally explored how chemical cues from a predator influence foraging behaviour and metabolic rate in females of the orb-web spider Argiope keyserlingi We found that female spiders restricted their foraging time budget when exposed to the predator cues from a praying mantid: they responded 11% and 17% quicker to a vibratory stimulus compared with control and non-predator cues, respectively, and spent less time handling the prey. Moreover, spiders were less likely to rebuild the web under predator cues. Female A. keyserlingi exposed to the praying mantid cue significantly elevated their metabolic rate compared with the control group. Our findings revealed short-term modifications over the 2 week trials in foraging behaviour and the physiology of female spiders in response to predator cues. This study suggests that under predator cues the spiders move quicker and this could be facilitated by elevation in metabolic rate. Reduced foraging activity and less frequent web repair/rebuilding would also reduce the spiders' exposure to praying mantid predators., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Spiders are known for producing specialized fibers. The radial orb-web, for example, contains tough silk used for the web frame and the capture spiral consists of elastic silk, able to stretch when prey impacts the web. In concert, silk proteins and web geometry affects the spider's ability to capture prey. Both factors have received considerable research attention, but next to no attention has been paid to the influence of fiber processing on web performance. Cribellate spiders produce a complex fiber alignment as their capture threads. With a temporally controlled spinneret movement, they connect different fibers at specific points to each other. One of the most complex capture threads is produced by the southern house spider, Kukulcania hibernalis (Filistatidae). In contrast to the so far characterized linear threads of other cribellate spiders, K. hibernalis spins capture threads in a zigzag pattern due to a slightly altered spinneret movement. The resulting more complex fiber alignment increased the thread's overall ability to restrain prey, probably by increasing the adhesion area as well as its extensibility. Kukulcania hibernalis' cribellate silk perfectly illustrates the impact of small behavioral differences on the thread assembly and, thus, of silk functionality.

Physical structures built by animals challenge our understanding of biological processes and inspire the development of smart materials and green architecture. It is thus indispensable to understand the drivers, constraints, and dynamics that lead to the emergence and modification of building behavior. Here, we demonstrate that spider web diversification repeatedly followed strikingly similar evolutionary trajectories, guided by physical constraints. We found that the evolution of suspended webs that intercept flying prey coincided with small changes in silk anchoring behavior with considerable effects on the robustness of web attachment. The use of nanofiber based capture threads (cribellate silk) conflicts with the behavioral enhancement of web attachment, and the repeated loss of this trait was frequently followed by physical improvements of web anchor structure. These findings suggest that the evolution of building behavior may be constrained by major physical traits limiting its role in rapid adaptation to a changing environment., (© 2019 The Author(s). Evolution © 2019 The Society for the Study of Evolution.)

Understanding how and why animal secretions vary in property has important biomimetic implications as desirable properties might covary. Spider major ampullate (MA) silk, for instance, is a secretion earmarked for biomimetic applications, but many of its properties vary among and between species across environments. Here, we tested the hypothesis that MA silk colour, protein structure and thermal properties covary when protein uptake is manipulated in the spider Trichonephila plumipes. We collected silk from adult female spiders maintained on a protein-fed or protein-deprived diet. Based on spectrophotometric quantifications, we classified half the silks as 'bee visible' and the other half 'bee invisible'. Wide angle X-ray diffraction and differential scanning calorimetry were then used to assess the silk's protein structure and thermal properties, respectively. We found that although protein structures and thermal properties varied across our treatments only the thermal properties covaried with colour. This ultimately suggests that protein structure alone is not responsible for MA silk thermal properties, nor does it affect silk colours. We speculate that similar ecological factors act on silk colour and thermal properties, which should be uncovered to inform biomimetic programmes.

The nature and size of attachments is a fundamental element of animal constructions. Presumably, these adhesive structures are plastically deployed to balance material investment and attachment strength. Here we studied plasticity in dragline anchorages of the golden orb web spider, Nephila plumipes. Specifically, we predict that spiders adjust the size and structure of dragline anchorages with load, i.e. spider mass. Mass was manipulated by attaching lead pieces to the spider's abdomen resulting in a 50 percent increase in mass. Loaded spiders spun larger but structurally similar thread anchorages than unloaded spiders. Thus, the spinning program that determines the overall anchor structure is highly stereotypic, and flexibility is introduced through varying the anchor size by increasing material investment. Our study showcases substrate attachments as suitable models to investigate the interplay between innate and changeable elements in the economy of building behaviours., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

Male reproductive performance can vary with condition, age and future reproductive opportunities. Web-building spiders are ideal models to examine the effects of senescence on fitness-related behaviours due to strong selection on male courtship to reduce pre-copulatory sexual cannibalism. Argiope keyserlingi spiders generate courtship vibrations, or 'shudders', that reduce female aggression. We found that male A. keyserlingi courtship slowed with chronological age. Older males took longer to travel across the courtship thread, and overall number of shudders increased. Males retained some ability to modulate courtship quality (shudder duration and number of rocks within each shudder) in response to female quality. A change in courtship performance over time, despite strong selection for repeatability, indicates that ageing in male A. keyserlingi may have direct impacts on reproductive performance.

Ostreid herpes virus causes serious disease in the Pacific oyster (Crassostrea gigas), but not in the Sydney Rock Oyster (Saccostrea glomerata). To investigate differences in disease progression, we injected oysters with double stranded RNA (dsRNA). dsRNA is known to mimic viral infection, and can evoke immune responses when Toll-like receptors detect the dsRNA, leading to the production of type 1 interferon and inflammation cytokines. The uptake and processing of dsRNA was tracked in gill and mantle tissue of Crassostrea gigas and Saccostrea glomerata after injection of fluorochrome labelled poly (I:C) dsRNA. The two species showed significant differences in tissue uptake and clearance, and differences in immune responses confirmed by real time PCR. These results showed that S. glomerata was more efficient in processing dsRNA than C. gigas, and that the gill tissue is an important site of dsRNA processing and response., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The evolution of a visual signal will be affected by signaler and receiver behavior, and by the physical properties of the environment where the signal is displayed. Crab spiders are typical sit-and-wait predators found in diverse ambush sites, such as tree bark, foliage, and flowers. Some of the flower-dweller species present a UV + -white visual lure that makes them conspicuous and attractive to their prey. We hypothesized that UV + -white coloration was associated with the evolution of a flower-dwelling habit. In addition, following up on results from a previous study we tested whether the UV + -white coloration evolved predominantly in flower-dwelling species occurring in Australia. We measured the reflectance of 1149 specimens from 66 species collected in Australia and Europe, reconstructed a crab spider phylogeny, and applied phylogenetic comparative methods to test our hypotheses. We found that the flower-dwelling habit evolved independently multiple times, and that this trait was correlated with the evolution of the UV + -white coloration. However, outside Australia non-flower-dwelling crab spiders also express a UV + -white coloration. Therefore, UV + -white reflectance is probably a recurring adaptation of some flower dwellers for attracting pollinators, although it may have other functions in non-flower dwellers, such as camouflage., (© 2017 The Author(s). Evolution © 2017 The Society for the Study of Evolution.)

Building behaviour in animals extends biological functions beyond bodies. Many studies have emphasized the role of behavioural programmes, physiology and extrinsic factors for the structure and function of buildings. Structure attachments associated with animal constructions offer yet unrealized research opportunities. Spiders build a variety of one- to three-dimensional structures from silk fibres. The evolution of economic web shapes as a key for ecological success in spiders has been related to the emergence of high performance silks and thread coating glues. However, the role of thread anchorages has been widely neglected in those models. Here, we show that orb-web (Araneidae) and hunting spiders (Sparassidae) use different silk application patterns that determine the structure and robustness of the joint in silk thread anchorages. Silk anchorages of orb-web spiders show a greater robustness against different loading situations, whereas the silk anchorages of hunting spiders have their highest pull-off resistance when loaded parallel to the substrate along the direction of dragline spinning. This suggests that the behavioural 'printing' of silk into attachment discs along with spinneret morphology was a prerequisite for the evolution of extended silk use in a three-dimensional space. This highlights the ecological role of attachments in the evolution of animal architectures., (© 2017 The Author(s).)

The non-consumptive effects of predation contribute to reduce preys' fitness. In this way, predation imposes a cost to animals, not only through direct consumption, but also as an energetic cost. One way used to estimate this cost in the past has been to measure the production of CO 2 to estimate the change in metabolic rate because of predation. It has been proposed that this change is mediated by the insect stress neurohormone octopamine. Here we study the change in metabolic rate of the black field cricket (Teleogryllus commodus), and how the production of CO 2 varies when a chemical cue from a sympatric predator is added. We hypothesised that after the addition of a predatory cue, the metabolic rate will increase. Moreover, since the pressure of predation is stronger on females, we propose that females will have a greater increase in the CO 2 produce as consequence of the added cues from the predator. Our results confirmed our first hypothesis, showing an almost two-fold increase in CO 2 when the predatory cue was added. However, males were the ones that showed a greater increase, in opposition to our second hypothesis. We put these results in the context of the escape theory and, in particular, the "landscape of fear" hypothesis. Also, because the timing between the increase of metabolic rate we measure here and the release of octopamine reported in previous studies do not match, we reject the idea that octopamine causes the increase in metabolism., (Copyright © 2017 Elsevier Inc. All rights reserved.)

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.

Mimicry complexes typically consist of multiple species that deter predators using similar anti-predatory signals. Mimics in these complexes are assumed to vary in their level of defence from highly defended through to moderately defended, or not defended at all. Here, we report a new multi-order mimicry complex that includes at least 140 different putative mimics from four arthropod orders including ants, wasps, bugs, tree hoppers and spiders. All members of this mimicry complex are characterised by a conspicuous golden body and an ant Gestalt , but vary substantially in their defensive traits. However, they were similarly effective at deterring predators - even mildly defended mimics were rarely eaten by a community of invertebrate and vertebrate predators both in the wild and during staged trials. We propose that despite the predominance of less defended mimics the three predatory guilds avoid the mimics because of the additive influence of the various defensive traits., Competing Interests: The authors declare that no competing interests exist.

The anchorage of structures is a crucial element of construction, both for humans and animals. Spiders use adhesive plaques to attach silk threads to substrates. Both biological and artificial adhesive structures usually have an optimal loading angle, and are prone to varying loading situations. Silk anchorages, however, must cope with loading in highly variable directions. Here we show that the detachment forces of thread anchorages of orb-web spiders are highly robust against pulling in different directions. This is gained by a two-step back-and-forth spinning pattern during the rapid production of the adhesive plaque, which shifts the thread insertion point towards the plaque centre and forms a flexible tree root-like network of branching fibres around the loading point. Using a morphometric approach and a tape-and-thread model we show that neither area, nor width of the plaque, but the shift of the loading point towards the plaque centre has the highest effect on pull-off resistance. This is explained by a circular propagation of the delamination crack with a low peeling angle. We further show that silken attachment discs are highly directional and adjusted to provide maximal performance in the upstream dragline. These results show that the way the glue is applied, crucially enhances the toughness of the anchorage without the need of additional material intake. This work is a starting point to study the evolution of tough and universal thread anchorages among spiders, and to develop bioinspired 'instant' anchorages of thread- and cable-like structures to a broad bandwidth of substrates., (© 2017 The Author(s).)

The use of ultraviolet (UV) cues for intra- and inter-specific communication is common in many animal species. Still, the role of UV signals under some predator-prey contexts, such as Batesian mimicry, is not clear. Batesian mimicry is a defensive strategy by which a palatable species (the mimic) resembles an unpalatable or noxious species (the model) to avoid predation. This strategy has evolved independently in many different taxa that are predated by species capable of UV perception. Moreover, there is considerable variation in how accurately Batesian mimics resemble their models across species. Our aim was to investigate how UV colour contributed to mimetic accuracy using several ant-mimicking spider species as a case study. We measured the reflectance spectrum (300-700 nm) for several species of mimics and models, and we tested whether they differ in visible and UV colour. We modelled whether two different predators could discriminate between mimics and models using colour information. We found that generally, ant-mimicking spiders differed significantly from their ant models in UV colour and that information from the visible range of light cannot be extrapolated into the UV. Our modelling suggested that wasps should be able to discriminate between mimics and models combining information from visible and the UV light, whereas birds may not discriminate between them. Thus, we show that UV colour can influence mimic accuracy and we discuss its potential role in Batesian mimicry. We conclude that colour, especially in the UV range, should be taken into account when measuring mimetic accuracy.

In groups of socially foraging animals, feeding behaviour may change with group size in response to varying cost-benefit trade-offs. Numerous studies have described group-size effects on group-average feeding behaviour, particularly emphasizing an increase in scrounging incidence for larger groups, where individuals (scroungers) feed from the food sources others (producers) discovered. However, individual variation in feeding behaviour remains unconsidered in the vast majority of these studies even though theoretical models predict individuals to specialize in feeding tactic and anticipate higher scrounger-type frequencies in larger groups. We combined group-level and individual-level analyses of group-size effects on social foraging in the subsocial spider Australomisidia ergandros Lending novel experimental support to model predictions, we found that individuals specialize in feeding tactic and that higher scrounging and lower producing incidence in larger groups were mediated through shifts in the ratio of feeding types. Further, feeding-type specialization was not explained by innate individual differences in hunting ability as all feeding types were equally efficient in prey capture when foraging alone. Context adaptivity of feeding behaviour might allow this subsocial species to succeed under varying socioecological conditions., (© 2016 The Author(s).)

The world in color presents a dazzling dimension of phenotypic variation. Biological interest in this variation has burgeoned, due to both increased means for quantifying spectral information and heightened appreciation for how animals view the world differently than humans. Effective study of color traits is challenged by how to best quantify visual perception in nonhuman species. This requires consideration of at least visual physiology but ultimately also the neural processes underlying perception. Our knowledge of color perception is founded largely on the principles gained from human psychophysics that have proven generalizable based on comparative studies in select animal models. Appreciation of these principles, their empirical foundation, and the reasonable limits to their applicability is crucial to reaching informed conclusions in color research. In this article, we seek a common intellectual basis for the study of color in nature. We first discuss the key perceptual principles, namely, retinal photoreception, sensory channels, opponent processing, color constancy, and receptor noise. We then draw on this basis to inform an analytical framework driven by the research question in relation to identifiable viewers and visual tasks of interest. Consideration of the limits to perceptual inference guides two primary decisions: first, whether a sensory-based approach is necessary and justified and, second, whether the visual task refers to perceptual distance or discriminability. We outline informed approaches in each situation and discuss key challenges for future progress, focusing particularly on how animals perceive color. Given that animal behavior serves as both the basic unit of psychophysics and the ultimate driver of color ecology/evolution, behavioral data are critical to reconciling knowledge across the schools of color research.

In alpine Australia, Orthoptera are abundant, dominant herbivores, important prey species, and hosts for parasites and parasitoids. Despite the central role of orthopterans in alpine ecosystems, the impact of parasites on orthopteran populations is under-explored. In this study we describe the relationship between parasite prevalence and host sex, body size and year of collection. We accessed an existing, preserved collection of 640 Kosciuscola tristis collected from across its range between 2007 and 2011. Upon dissection we collected juvenile parasites and used molecular tools to identify them to three families (Nematoda; Mermithidae, and Arthropoda: Diptera: Tachinidae and Sarcophagidae). The prevalence of nematodes ranged from 3.5% to 25.0% and dipterans from 2.4% to 20.0%. Contrary to predictions, we found no associations between parasite prevalence and grasshopper sex or size. Although there was an association between prevalence of both nematodes and dipterans with year of collection, this is likely driven by a small sample size in the first year. Our results provide a foundation for future studies into parasite prevalence within the alpine environment and the abiotic factors that might influence these associations.

The sterile male technique is a common method to assign paternity, widely adopted due to its relative simplicity and low cost. Male sterility is induced by exposure to sub lethal doses of chemosterilants or irradiation, the dosage of which has to be calibrated for every species to provide successful male sterilisation, without affecting male physiology and behaviour. While the physiological effects of sterilisation are usually assessed for each study, the behavioural ones are rarely analysed in detail. Using the orb web spider Argiope keyserlingi as a model we first tested (1) the validity of the thread assay, which simulates male courtship behaviour in a standardised context, as a proxy representing courtship on a female web. We then investigated (2) the effectiveness of male sterilisation via irradiation and (3) its consequences on male courtship behaviour. Our results validate the thread assay and the sterile male technique as legitimate tools for the study of male courtship behaviour and fertilisation success. We show that these techniques are time and cost effective and reduce undesirable variation, thereby creating opportunities to study and understand the mechanisms underlying sexual selection., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The mechanisms and functions of reversible colour change in arthropods are highly diverse despite, or perhaps due to, the presence of an exoskeleton. Physiological colour changes, which have been recorded in 90 arthropod species, are rapid and are the result of changes in the positioning of microstructures or pigments, or in the refractive index of layers in the integument. By contrast, morphological colour changes, documented in 31 species, involve the anabolism or catabolism of components (e.g. pigments) directly related to the observable colour. In this review we highlight the diversity of mechanisms by which reversible colour change occurs and the evolutionary context and diversity of arthropod taxa in which it has been observed. Further, we discuss the functions of reversible colour change so far proposed, review the limited behavioural and ecological data, and argue that the field requires phylogenetically controlled approaches to understanding the evolution of reversible colour change. Finally, we encourage biologists to explore new model systems for colour change and to engage scientists from other disciplines; continued cross-disciplinary collaboration is the most promising approach to this nexus of biology, physics, and chemistry., (© 2014 The Authors. Biological Reviews © 2014 Cambridge Philosophical Society.)

The diversity, variability, and apparent rapid evolution of animal genitalia are a vivid focus of research in evolutionary biology, and studies exploring genitalia have dramatically increased over the past decade. These studies, however, exhibit a strong male bias, which has worsened since 2000, despite the fact that this bias has been explicitly pointed out in the past. Early critics argued that previous investigators too often considered only males and their genitalia, while overlooking female genitalia or physiology. Our analysis of the literature shows that overall this male bias has worsened with time. The degree of bias is not consistent between subdisciplines: studies of the lock-and-key hypothesis have been the most male focused, while studies of cryptic female choice usually consider both sexes. The degree of bias also differed across taxonomic groups, but did not associate with the ease of study of male and female genital characteristics. We argue that the persisting male bias in this field cannot solely be explained by anatomical sex differences influencing accessibility. Rather the bias reflects enduring assumptions about the dominant role of males in sex, and invariant female genitalia. New research highlights how rapidly female genital traits can evolve, and how complex coevolutionary dynamics between males and females can shape genital structures. We argue that understanding genital evolution is hampered by an outdated single-sex bias., Competing Interests: The authors have declared that no competing interests exist.

In brood-caring species, family members are faced with a conflict over resource distribution. While parents are selected to adapt the amount of care according to their offspring's needs, offspring might be selected to demand more care than optimal for parents. Recent studies on birds have shown that the social network structure of offspring affects the amount of care and thus the fitness of families. Such a network structure of repeated interactions is probably influenced by within-brood relatedness. We experimentally manipulated the group composition in a brood-caring spider to test how the presence of unrelated spiderlings affects the dynamics between female and brood as well as within broods. Broods consisting of siblings grew better and had a lower mortality compared with mixed broods, no matter whether the caring female was a genetic or foster mother. Interestingly, we found that foster mothers lost weight when caring for sibling broods, whereas females caring for mixed broods gained weight. This indicates that females may be willing to share more prey when the brood contains exclusively siblings even if the entire brood is unrelated to the female. Resource distribution may thus be negotiated by offspring dynamics that could have a signalling function to females.

Mimicry has evolved in contexts such as camouflage, predator deterrence, luring of prey, and pollinator attraction. Mimicry of flowers has until now been demonstrated only in angiosperms, yet it has been hypothesized that the Malaysian orchid mantis Hymenopus coronatus mimics a flower to attract pollinators as prey. Despite the popularity of this charismatic insect, this long-discussed hypothesis has never been experimentally investigated. We found that, as predicted for mimicry, the color of H. coronatus is indistinguishable from the color of sympatric flowers for hymenopteran pollinators. Field experiments show that isolated mantises attract wild pollinators at a rate even higher than flowers and capture these pollinators as prey items. After more than a century of conjecture, we provide the first experimental evidence of pollinator deception in the orchid mantis and the first description of a unique predatory strategy that has not been documented in any other animal species.

During courtship, individuals transfer information about identity, mating status and quality. However, male web-building spiders face a significant problem: how to begin courting female spiders without being mistaken for prey? Male Argiope spiders generate distinctive courtship vibrations (shudders) when entering a female's web. We tested whether courtship shudders delay female predatory behaviour, even when live prey is present in the web. We presented a live cricket to females during playbacks of shudder vibrations, or white noise, and compared female responses to a control in which we presented a live cricket with no playback vibrations. Females were much slower to respond to crickets during playback of shudder vibrations. Shudder vibrations also delayed female predatory behaviour in a related spider species, showing that these vibrations do not simply function for species identity. These results suggest that male web-building spiders employ a phylogenetically conserved vibratory signal to ameliorate the risk of pre-copulatory cannibalism.

Theory suggests that aposematism, specifically the learned avoidance of unprofitable prey via memorable color patterns, should result in selection for pattern uniformity. However, many examples to the contrary are seen in nature. Conversely, honest sexual signals are likely to exhibit greater variation because they reflect underlying variation in mate quality. Here we aim to characterize and quantify the mechanistic causes of color in Tectocoris diophthalmus [corrected] to shed light on the costs of color production, and thus the potential information content of its color signals. We use Tectocoris diophthalmus [corrected] because it is a weakly-defended stinkbug, and presents elements that have classically been studied in the context of aposematism (red coloring), and sexual selection (sexual dichromatism and iridescent coloring). Pigment analysis reveals that variation in orange coloration is due to the amount of erythropterin pigment, stored in intracellular granules. This pigment is common in Heteroptera, and as an endogenously produced excretory byproduct is unlikely to reflect mate quality or variation in unprofitability of the bug. Electron microscopy reveals the iridescent patches are caused by an epicuticular multilayer reflector, and the hue and patch size are directly related to the layer widths and extent of coverage of this layering. Furthermore, we identified melanin as an essential component of the multilayer reflector system; therefore, the quality of the iridescent patches may be affected by aspects of rearing environment and immunocompetence. We posit that T. diophthalmus [corrected] has co-opted the melanic patches of a 'typical' red and black aposematic signal, transforming it into a complex and variable iridescent signal that may enhance its capacity to display individual quality.

Web-building spiders are important models for sexual selection. While our understanding of post-copulatory mechanisms including sperm competition and cryptic female choice is considerable, our knowledge of courtship and how it influences male and female mating decisions is still extremely poor. Here, we provide the first comprehensive description of male courtship behaviour and vibrations generated in the web by the orb-web spider, Argiope keyserlingi - a recognised model species. We identified three main elements of male courtship: shudders, abdominal wags and mating thread dances (including both plucks and bounces). The vibrations generated by these behaviours are described in detail. Male shuddering behaviour appears to have a strong influence on female latency to mate acceptance, with males that shudder at high rates without compromising shudder duration being preferred. Shuddering behaviour may also mediate female aggressive behaviour, with males that generate long shudders less likely to be cannibalised after copulation. Male abdominal wagging behaviour, however, appears to have only limited influence on female mating decisions. This study provides avenues for future work that synthesises pre- and post-copulatory mechanisms in web-building spiders to generate an all-encompassing model of how sexual selection operates.

Sit-and-wait predators have evolved several traits that increase the probability of encountering prey, including lures that attract prey. Although most crab spiders (Thomisidae) are known by their ability to change colour in order to match the background, a few use a different strategy. They are UV-reflective, creating a colour contrast against UV-absorbing flowers that is attractive for pollinators. The nature of the relationship between colour contrast and foraging success is unknown, as is how spiders trade off the potential costs and benefits of strong colour contrast. Therefore, this study investigated the relationship between spider colouration, foraging success and background colouration in a crab spider species known to lure pollinators via UV reflectance (Thomisus spectabilis). Field data revealed that spider body condition - a proxy of past foraging success - is positively related to overall colour contrast. We experimentally tested the effect of satiation and background colour on spider colour change. Throughout the experiment, spiders changed their colour contrast regardless of their food intake, suggesting that colour contrast and the UV component contributing to overall contrast are not caused by spider condition. Although spiders responded to different backgrounds by subtly changing their body colour, this did not result in colour matching. We believe that the observed variation in colour contrast and hence conspicuousness in the field, coupled with the spiders' reaction to our manipulation, could be the result of plasticity in response to prey.