No sex difference in preen oil chemical composition during incubation in Kentish plovers.

Preen oil, the secretion from the uropygial gland of birds, may have a specific function in incubation. Consistent with this, during incubation, the chemical composition of preen oil is more likely to differ between sexes in species where only one sex incubates than in species where both sexes incubate. In this study, we tested the generality of this apparent difference, by investigating sex differences in the preen oil composition of a shorebird species, the Kentish plover (Anarhynchus, formerly Charadrius, alexandrinus). As both sexes incubate in this species, we predicted the absence of sex differences in preen oil composition during incubation. In the field, we sampled preen oil from nine females and 11 males during incubation, which we analysed with gas chromatography-mass spectrometry (GC-MS). Consistent with predictions, we found no sex difference in preen oil composition, neither in beta diversity (Bray-Curtis dissimilarities) nor in alpha diversity (Shannon index and number of substances). Based on these results, we cannot conclude whether preen oil has a function during incubation in Kentish plovers. Still, we discuss hypothetical roles, such as olfactory crypsis, protection against ectoparasites or olfactory intraspecific communication, which remain to be tested.

Interactions between color and gloss in iridescent camouflage.

Iridescence is a taxonomically widespread form of structural coloration that produces often intense hues that change with the angle of viewing. Its role as a signal has been investigated in multiple species, but recently, and counter-intuitively, it has been shown that it can function as camouflage. However, the property of iridescence that reduces detectability is, as yet, unclear. As viewing angle changes, iridescent objects change not only in hue but also in intensity, and many iridescent animals are also shiny or glossy; these "specular reflections," both from the target and background, have been implicated in crypsis. Here, we present a field experiment with natural avian predators that separate the relative contributions of color and gloss to the "survival" of iridescent and non-iridescent beetle-like targets. Consistent with previous research, we found that iridescent coloration, and high gloss of the leaves on which targets were placed, enhance survival. However, glossy targets survived less well than matt. We interpret the results in terms of signal-to-noise ratio: specular reflections from the background reduce detectability by increasing visual noise. While a specular reflection from the target attracts attention, a changeable color reduces the signal because, we suggest, normally, the color of an object is a stable feature for detection and identification.

Pattern variation is linked to anti-predator coloration in butterfly larvae.

Prey animals typically try to avoid being detected and/or advertise to would-be predators that they should be avoided. Both anti-predator strategies primarily rely on colour to succeed, but the specific patterning used is also important. While the role of patterning in camouflage is relatively clear, the design features of aposematic patterns are less well understood. Here, we use a comparative approach to investigate how pattern use varies across a phylogeny of 268 species of cryptic and aposematic butterfly larvae, which also vary in social behaviour. We find that longitudinal stripes are used more frequently by cryptic larvae, and that patterns putatively linked to crypsis are more likely to be used by solitary larvae. By contrast, aposematic larvae are more likely to use horizontal bands and spots, but we find no differences in the use of individual pattern elements between solitary and gregarious aposematic species. However, solitary aposematic larvae are more likely to display multiple pattern elements, whereas those with no pattern are more likely to be gregarious. Our study advances our understanding of how pattern variation, coloration and social behaviour covary across lepidopteran larvae, and highlights new questions about how patterning affects larval detectability and predator responses to aposematic prey.

Warning Coloration, Body Size, and the Evolution of Gregarious Behavior in Butterfly Larvae.

AbstractMany species gain antipredator benefits by combining gregarious behavior with warning coloration, yet there is debate over which trait evolves first and which is the secondary adaptive enhancement. Body size can also influence how predators receive aposematic signals and potentially constrain the evolution of gregarious behavior. To our knowledge, the causative links between the evolution of gregariousness, aposematism, and larger body sizes have not been fully resolved. Here, using the most recently resolved butterfly phylogeny and an extensive new dataset of larval traits, we reveal the evolutionary interactions between important traits linked to larval gregariousness. We show that larval gregariousness has arisen many times across butterflies, and aposematism is a likely prerequisite for gregariousness to evolve. We also find that body size may be an important factor for determining the coloration of solitary, but not gregarious, larvae. Additionally, by exposing artificial larvae to wild avian predation, we show that undefended, cryptic larvae are heavily predated when aggregated but benefit from solitariness, whereas the reverse is true for aposematic prey. Our data reinforce the importance of aposematism for gregarious larval survival while identifying new questions about the roles of body size and toxicity in the evolution of grouping behavior.

Remating opportunities and low costs underlie maternal desertion.

Parental care can enhance offspring survival but may impose significant costs to parents. The costs and benefits of care are key to understanding patterns of parental care, where parents can benefit by having their partner increase investment in care, while reducing their own effort. However, investigating the costs and benefits of parental care in wild populations is challenging. Here we use highly detailed behavioral observations in families of a small shorebird, where one parent frequently deserts its offspring, to explore the potential costs and benefits of desertion in a wild population. We first show that females desert their broods more frequently than males. Second, we investigate the benefits of this frequent female desertion in terms of additional mating opportunities, and the costs of desertion to females in terms of the growth and survival of deserted offspring. Our results indicate that female desertion is favored by a combination of remating benefits and a lack of costs to brood growth and survival, as abandoned male parents continue to provide care after desertion. Our results shed light on the costs and benefits underlying natural desertion strategies and suggest that female desertion is a fine-tuned behavior that responds to seasonally changing benefits of desertion.

Olfactory camouflage and communication in birds.

Smell is a sensory modality that is rarely considered in birds, but evidence is mounting that olfaction is an important aspect of avian behaviour and ecology. The uropygial gland produces an odoriferous secretion (preen oil) that can differ seasonally and between the sexes. These differences are hypothesized to function in olfactory camouflage, i.e. minimizing detection by nest predators (olfactory crypsis hypothesis), and/or intraspecific olfactory communication, particularly during breeding (sex semiochemical hypothesis). However, evidence for seasonal and sex differences in preen oil is mixed, with some studies finding differences and others not, and direct evidence for the putative function(s) of seasonal variation and sex differences in preen oil remains limited. We conducted a systematic review of the evidence for such changes in preen oil chemical composition, finding seasonal differences in 95% of species (57/60 species in 35 studies) and sex differences in 47% of species (28/59 species in 46 studies). We then conducted phylogenetic comparative analyses using data from 59 bird species to evaluate evidence for both the olfactory crypsis and sex semiochemical hypotheses. Seasonal differences were more likely in the incubating than non-incubating sex in ground-nesting species, but were equally likely regardless of incubation strategy in non-ground-nesting species. This result supports the olfactory crypsis hypothesis, if ground nesters are more vulnerable to olfactorily searching predators than non-ground nesters. Sex differences were more likely in species with uniparental than biparental incubation and during breeding than non-breeding, consistent with both the olfactory crypsis and sex semiochemical hypotheses. At present, the data do not allow us to disentangle these two hypotheses, but we provide recommendations that will enable researchers to do so.

Beetle iridescence induces an avoidance response in naïve avian predators.

It has recently been found that iridescence, a taxonomically widespread form of animal coloration defined by a change in hue with viewing angle, can act as a highly effective form of camouflage. However, little is known about whether iridescence can confer a survival benefit to prey postdetection and, if so, which optical properties of iridescent prey are important for this putative protective function. Here, we tested the effects of both iridescence and surface gloss (i.e. specular reflection) on the attack behaviour of prey-naïve avian predators. Using real and artificial jewel beetle, Sternocera aequisignata, wing cases, we found that iridescence provides initial protection against avian predation by significantly reducing the willingness to attack. Importantly, we found that the main factor explaining this aversion is iridescence, not multiple colours per se, with surface gloss also having an independent effect. Our results are important because they demonstrate that even when prey are presented up close and against a mismatching background, iridescence may confer a survival benefit by inducing hesitation or even, as sometimes observed, an aversion response in attacking birds. Furthermore, this means that even postdetection, prey do not necessarily need to have secondary defences such as sharp spines or toxins for iridescence to have a protective effect. Taken together, our results suggest that reduced avian predation could facilitate the initial evolution of iridescence in many species of insects and that it is the defining feature of iridescence, its colour changeability, that is important for this effect.

Background complexity can mitigate poor camouflage.

Avoiding detection through camouflage is often key to survival. However, an animal's appearance is not the only factor affecting conspicuousness: background complexity also alters detectability. This has been experimentally demonstrated for both artificially patterned backgrounds in the laboratory and natural backgrounds in the wild, but only for targets that already match the background well. Do habitats of high visual complexity provide concealment to even relatively poorly camouflaged animals? Using artificial prey which differed in their degrees of background matching to tree bark, we were able to determine their survival, under bird predation, with respect to the natural complexity of the background. The latter was quantified using low-level vision metrics of feature congestion (or 'visual clutter') adapted for bird vision. Higher background orientation clutter (edges with varying orientation) reduced the detectability of all but the poorest background-matching camouflaged treatments; higher background luminance clutter (varying achromatic lightness) reduced average mortality for all treatments. Our results suggest that poorer camouflage can be mitigated by more complex backgrounds, with implications for both camouflage evolution and habitat preferences.

Birds learn to avoid aposematic prey by using the appearance of host plants.

The conspicuous warning signal of aposematic animals is learned by their predators, and the resulting avoidance benefits both parties.1-4 Given evidence that birds can distinguish the profitability of prey from the environmental context in which they appear,5 aposematic insects' host plants might also provide an important cue to foraging predators.6 The aposematic cinnabar moth (Tyria jacobaeae) larva is a specialist on its ragwort (Senecio spp.) host plant,7 presenting a consistent environment with which it could be reliably associated. Additionally, ragwort's defensive toxins prevent non-specialist, profitable insects from feeding on it.8 Thus, avian predators may recognize cues from ragwort, most likely its conspicuous yellow flowers,9,10 and use this information to avoid cinnabars. To test this hypothesis, we exposed artificial cinnabar and non-signaling "caterpillar" targets to wild avian predation by presenting them on ragwort and non-toxic host plants. We also manipulated the presence or absence of ragwort flowers on hosts. In doing so, we show that both targets are better protected on the cinnabar's natural ragwort host and that birds use ragwort's distinctive yellow flowers as the cue to avoidance. Additionally, we found that naive predators do not make prey host foraging distinctions, indicating that this avoidance behavior is learned through experience. Our findings are among the first to suggest that a host plant's features act as an extended phenotype that signals the toxicity of the prey that live on it. This prey-host relationship may facilitate the initial evolution of toxicity in non-signaling prey, but also inhibit the evolution of aposematic signals themselves. VIDEO ABSTRACT.

Nature calls: intelligence and natural foraging style predict poor welfare in captive parrots.

Understanding why some species thrive in captivity, while others struggle to adjust, can suggest new ways to improve animal care. Approximately half of all Psittaciformes, a highly threatened order, live in zoos, breeding centres and private homes. Here, some species are prone to behavioural and reproductive problems that raise conservation and ethical concerns. To identify risk factors, we analysed data on hatching rates in breeding centres (115 species, 10 255 pairs) and stereotypic behaviour (SB) in private homes (50 species, 1378 individuals), using phylogenetic comparative methods (PCMs). Small captive population sizes predicted low hatch rates, potentially due to genetic bottlenecks, inbreeding and low availability of compatible mates. Species naturally reliant on diets requiring substantial handling were most prone to feather-damaging behaviours (e.g. self-plucking), indicating inadequacies in the composition or presentation of feed (often highly processed). Parrot species with relatively large brains were most prone to oral and whole-body SB: the first empirical evidence that intelligence can confer poor captive welfare. Together, results suggest that more naturalistic diets would improve welfare, and that intelligent psittacines need increased cognitive stimulation. These findings should help improve captive parrot care and inspire further PCM research to understand species differences in responses to captivity.

Colour pattern variation forms local background matching camouflage in a leaf-mimicking toad.

Optimal camouflage can, in principle, be relatively easily achieved in simple, homogeneous, environments where backgrounds always have the same colour, brightness and patterning. Natural environments are, however, rarely homogenous, and species often find themselves viewed against varied backgrounds where the task of concealment is more challenging. One result of variable backgrounds is the evolution of intraspecific phenotypic variation which may either be generalized, with multiple similarly cryptic patterns, or specialized, with each discrete colour form maximizing concealment against a single component of the background. We investigated the role of phenotypic variation in a highly variable population of the Neotropical toad Rhinella margaritifera using visual modelling and a computer-based detection task. We found that phenotypic variation was not divided into discrete colour morphs, and all toads were well camouflaged against the forest floor. However, although the whole population may appear to consist of random samples from the background, the toads were a particularly close match to the leaf litter, suggesting that they masquerade as dead leaves, which are themselves variable. Furthermore, rather than each colour form being equally effective against a single background, each toad was specialized towards its own particular local surroundings, as suggested by a specialist strategy. Taken together, these data highlight the importance of background matching to a nominally masquerading species, as well as how habitat heterogeneity at multiple spatial scales may affect the evolution of camouflage and phenotypic variation.

The Camouflage Machine: Optimizing protective coloration using deep learning with genetic algorithms.

Evolutionary biologists frequently wish to measure the fitness of alternative phenotypes using behavioral experiments. However, many phenotypes are complex. One example is coloration: camouflage aims to make detection harder, while conspicuous signals (e.g., for warning or mate attraction) require the opposite. Identifying the hardest and easiest to find patterns is essential for understanding the evolutionary forces that shape protective coloration, but the parameter space of potential patterns (colored visual textures) is vast, limiting previous empirical studies to a narrow range of phenotypes. Here, we demonstrate how deep learning combined with genetic algorithms can be used to augment behavioral experiments, identifying both the best camouflage and the most conspicuous signal(s) from an arbitrarily vast array of patterns. To show the generality of our approach, we do so for both trichromatic (e.g., human) and dichromatic (e.g., typical mammalian) visual systems, in two different habitats. The patterns identified were validated using human participants; those identified as the best for camouflage were significantly harder to find than a tried-and-tested military design, while those identified as most conspicuous were significantly easier to find than other patterns. More generally, our method, dubbed the "Camouflage Machine," will be a useful tool for identifying the optimal phenotype in high dimensional state spaces.

Large Lemurs: Ecological, Demographic and Environmental Risk Factors for Weight Gain in Captivity.

Excessive body mass, i.e., being overweight or obese, is a health concern associated with issues such as reduced fertility and lifespan. Some lemur species are prone to extreme weight gain in captivity, yet others are not. To better understand species- and individual-level effects on susceptibility to captive weight gain, we use two complementary methods: phylogenetic comparative methods to examine ecological explanations for susceptibility to weight gain across species, and epidemiological approaches to examine demographic and environment effects within species. Data on body masses and living conditions were collected using a survey, yielding useable data on 675 lemurs representing 13 species from 96 collections worldwide. Data on species-typical wild ecology for comparative analyses came from published literature and climate databases. We uncovered one potential ecological risk factor: species adapted to greater wild food resource unpredictability tended to be more prone to weight gain. Our epidemiological analyses on the four best-sampled species revealed four demographic and one environmental risk factors, e.g., for males, being housed with only fixed climbing structures. We make practical recommendations to help address weight concerns, and describe future research including ways to validate the proxy we used to infer body condition.

The ARRIVE guidelines 2.0: Updated guidelines for reporting animal research.

Reproducible science requires transparent reporting. The ARRIVE guidelines (Animal Research: Reporting of In Vivo Experiments) were originally developed in 2010 to improve the reporting of animal research. They consist of a checklist of information to include in publications describing in vivo experiments to enable others to scrutinise the work adequately, evaluate its methodological rigour, and reproduce the methods and results. Despite considerable levels of endorsement by funders and journals over the years, adherence to the guidelines has been inconsistent, and the anticipated improvements in the quality of reporting in animal research publications have not been achieved. Here, we introduce ARRIVE 2.0. The guidelines have been updated and information reorganised to facilitate their use in practice. We used a Delphi exercise to prioritise and divide the items of the guidelines into 2 sets, the "ARRIVE Essential 10," which constitutes the minimum requirement, and the "Recommended Set," which describes the research context. This division facilitates improved reporting of animal research by supporting a stepwise approach to implementation. This helps journal editors and reviewers verify that the most important items are being reported in manuscripts. We have also developed the accompanying Explanation and Elaboration (E&E) document, which serves (1) to explain the rationale behind each item in the guidelines, (2) to clarify key concepts, and (3) to provide illustrative examples. We aim, through these changes, to help ensure that researchers, reviewers, and journal editors are better equipped to improve the rigour and transparency of the scientific process and thus reproducibility.

The ARRIVE guidelines 2.0: Updated guidelines for reporting animal research.

Reproducible science requires transparent reporting. The ARRIVE guidelines (Animal Research: Reporting of In Vivo Experiments) were originally developed in 2010 to improve the reporting of animal research. They consist of a checklist of information to include in publications describing in vivo experiments to enable others to scrutinise the work adequately, evaluate its methodological rigour, and reproduce the methods and results. Despite considerable levels of endorsement by funders and journals over the years, adherence to the guidelines has been inconsistent, and the anticipated improvements in the quality of reporting in animal research publications have not been achieved. Here, we introduce ARRIVE 2.0. The guidelines have been updated and information reorganised to facilitate their use in practice. We used a Delphi exercise to prioritise and divide the items of the guidelines into 2 sets, the "ARRIVE Essential 10," which constitutes the minimum requirement, and the "Recommended Set," which describes the research context. This division facilitates improved reporting of animal research by supporting a stepwise approach to implementation. This helps journal editors and reviewers verify that the most important items are being reported in manuscripts. We have also developed the accompanying Explanation and Elaboration document, which serves (1) to explain the rationale behind each item in the guidelines, (2) to clarify key concepts, and (3) to provide illustrative examples. We aim, through these changes, to help ensure that researchers, reviewers, and journal editors are better equipped to improve the rigour and transparency of the scientific process and thus reproducibility.

The ARRIVE guidelines 2.0: Updated guidelines for reporting animal research.

Reproducible science requires transparent reporting. The ARRIVE guidelines (Animal Research: Reporting of In Vivo Experiments) were originally developed in 2010 to improve the reporting of animal research. They consist of a checklist of information to include in publications describing in vivo experiments to enable others to scrutinise the work adequately, evaluate its methodological rigour, and reproduce the methods and results. Despite considerable levels of endorsement by funders and journals over the years, adherence to the guidelines has been inconsistent, and the anticipated improvements in the quality of reporting in animal research publications have not been achieved. Here, we introduce ARRIVE 2.0. The guidelines have been updated and information reorganised to facilitate their use in practice. We used a Delphi exercise to prioritise and divide the items of the guidelines into 2 sets, the "ARRIVE Essential 10," which constitutes the minimum requirement, and the "Recommended Set," which describes the research context. This division facilitates improved reporting of animal research by supporting a stepwise approach to implementation. This helps journal editors and reviewers verify that the most important items are being reported in manuscripts. We have also developed the accompanying Explanation and Elaboration (E&E) document, which serves (1) to explain the rationale behind each item in the guidelines, (2) to clarify key concepts, and (3) to provide illustrative examples. We aim, through these changes, to help ensure that researchers, reviewers, and journal editors are better equipped to improve the rigour and transparency of the scientific process and thus reproducibility.

Reporting animal research: Explanation and elaboration for the ARRIVE guidelines 2.0.

Improving the reproducibility of biomedical research is a major challenge. Transparent and accurate reporting is vital to this process; it allows readers to assess the reliability of the findings and repeat or build upon the work of other researchers. The ARRIVE guidelines (Animal Research: Reporting In Vivo Experiments) were developed in 2010 to help authors and journals identify the minimum information necessary to report in publications describing in vivo experiments. Despite widespread endorsement by the scientific community, the impact of ARRIVE on the transparency of reporting in animal research publications has been limited. We have revised the ARRIVE guidelines to update them and facilitate their use in practice. The revised guidelines are published alongside this paper. This explanation and elaboration document was developed as part of the revision. It provides further information about each of the 21 items in ARRIVE 2.0, including the rationale and supporting evidence for their inclusion in the guidelines, elaboration of details to report, and examples of good reporting from the published literature. This document also covers advice and best practice in the design and conduct of animal studies to support researchers in improving standards from the start of the experimental design process through to publication.

The ARRIVE guidelines 2.0: Updated guidelines for reporting animal research.

Reproducible science requires transparent reporting. The ARRIVE guidelines (Animal Research: Reporting of In Vivo Experiments) were originally developed in 2010 to improve the reporting of animal research. They consist of a checklist of information to include in publications describing in vivo experiments to enable others to scrutinise the work adequately, evaluate its methodological rigour, and reproduce the methods and results. Despite considerable levels of endorsement by funders and journals over the years, adherence to the guidelines has been inconsistent, and the anticipated improvements in the quality of reporting in animal research publications have not been achieved. Here, we introduce ARRIVE 2.0. The guidelines have been updated and information reorganised to facilitate their use in practice. We used a Delphi exercise to prioritise and divide the items of the guidelines into 2 sets, the "ARRIVE Essential 10," which constitutes the minimum requirement, and the "Recommended Set," which describes the research context. This division facilitates improved reporting of animal research by supporting a stepwise approach to implementation. This helps journal editors and reviewers verify that the most important items are being reported in manuscripts. We have also developed the accompanying Explanation and Elaboration document, which serves (1) to explain the rationale behind each item in the guidelines, (2) to clarify key concepts, and (3) to provide illustrative examples. We aim, through these changes, to help ensure that researchers, reviewers, and journal editors are better equipped to improve the rigour and transparency of the scientific process and thus reproducibility.

The ARRIVE guidelines 2.0: updated guidelines for reporting animal research.

Reproducible science requires transparent reporting. The ARRIVE guidelines (Animal Research: Reporting of In Vivo Experiments) were originally developed in 2010 to improve the reporting of animal research. They consist of a checklist of information to include in publications describing in vivo experiments to enable others to scrutinise the work adequately, evaluate its methodological rigour, and reproduce the methods and results. Despite considerable levels of endorsement by funders and journals over the years, adherence to the guidelines has been inconsistent, and the anticipated improvements in the quality of reporting in animal research publications have not been achieved. Here, we introduce ARRIVE 2.0. The guidelines have been updated and information reorganised to facilitate their use in practice. We used a Delphi exercise to prioritise and divide the items of the guidelines into 2 sets, the 'ARRIVE Essential 10,' which constitutes the minimum requirement, and the 'Recommended Set,' which describes the research context. This division facilitates improved reporting of animal research by supporting a stepwise approach to implementation. This helps journal editors and reviewers verify that the most important items are being reported in manuscripts. We have also developed the accompanying Explanation and Elaboration document, which serves (1) to explain the rationale behind each item in the guidelines, (2) to clarify key concepts, and (3) to provide illustrative examples. We aim, through these changes, to help ensure that researchers, reviewers, and journal editors are better equipped to improve the rigour and transparency of the scientific process and thus reproducibility.

The ARRIVE guidelines 2.0: Updated guidelines for reporting animal research.

Reproducible science requires transparent reporting. The ARRIVE guidelines (Animal Research: Reporting of In Vivo Experiments) were originally developed in 2010 to improve the reporting of animal research. They consist of a checklist of information to include in publications describing in vivo experiments to enable others to scrutinise the work adequately, evaluate its methodological rigour, and reproduce the methods and results. Despite considerable levels of endorsement by funders and journals over the years, adherence to the guidelines has been inconsistent, and the anticipated improvements in the quality of reporting in animal research publications have not been achieved. Here, we introduce ARRIVE 2.0. The guidelines have been updated and information reorganised to facilitate their use in practice. We used a Delphi exercise to prioritise and divide the items of the guidelines into 2 sets, the "ARRIVE Essential 10," which constitutes the minimum requirement, and the "Recommended Set," which describes the research context. This division facilitates improved reporting of animal research by supporting a stepwise approach to implementation. This helps journal editors and reviewers verify that the most important items are being reported in manuscripts. We have also developed the accompanying Explanation and Elaboration document, which serves (1) to explain the rationale behind each item in the guidelines, (2) to clarify key concepts, and (3) to provide illustrative examples. We aim, through these changes, to help ensure that researchers, reviewers, and journal editors are better equipped to improve the rigour and transparency of the scientific process and thus reproducibility.