The relationship between the body and air temperature in a terrestrial ectotherm.

Ectotherms make up the majority of terrestrial biodiversity, so it is important to understand their potential responses to climate change. Often, models aiming to achieve this understanding correlate species distributions with ambient air temperature. However, this assumes a constant relationship between the air temperature and body temperature, which determines an ectotherm's thermal performance. To test this assumption, we develop and validate a method for retrospective estimation of ectotherm body temperature using heat exchange equations. We apply the model to predict the body temperature of wild field crickets (Gryllus campestris) in Northern Spain for 1985-2019 and compare these values to air temperature. We show that while air temperature impacts ectotherm body temperature, it captures only a fraction of its thermal experience. Solar radiation can increase the body temperature by more than 20°C above air temperature with implications for physiology and behaviour. The effect of solar radiation on body temperature is particularly important given that climate change will alter cloud cover. Our study shows that the impacts of climate change on species cannot be assumed to be proportional only to changing air temperature. More reliable models of future species distributions require mechanistic links between environmental conditions and thermal ecophysiologies of species.

Environment and mate attractiveness in a wild insect.

The role of female choice in sexual selection is well established, including the recognition that females choose their mates based on multiple cues. These cues may include intrinsic aspects of a male's phenotype as well as aspects of the environment associated with the male. The role of the spatial location of a potential mate has been well studied in territorial vertebrates. However, despite their role as laboratory models for studies of sexual selection, the potential for insects to choose their mates on the basis of location has scarcely been studied. We studied a natural population of individually tagged crickets (Gryllus campestris) in a meadow in Northern Spain. Adults typically move between burrows every few days, allowing us to examine how pairing success of males can be predicted by the burrow they occupy, independent of their own characteristics. We observed the entirety of ten independent breeding seasons to provide replication and to determine whether the relative importance of these factors is stable across years. We find that both male ID and the ID his burrow affect the likelihood that he is paired with a female, but the burrow has a consistently greater influence. Furthermore, the two factors interact: the relative attractiveness of an individual male depends on which burrow he occupies. Our finding demonstrates a close interaction between naturally and sexually selected traits. It also demonstrates that mate choice studies may benefit from considering not only obvious secondary sexual traits, but also more cryptic traits such as microhabitat choice.

Anthropogenic noise may impair the mating behaviour of the Shore Crab Carcinus Maenas.

Anthropogenic noise is a recent addition to the list of human-made threats to the environment, with potential and established negative impacts on a wide range of animals. Despite their economic and ecological significance, few studies have considered the impact of anthropogenic noise on crustaceans, though past studies have shown that it can cause significant effects to crustacean physiology, anatomy, and behaviour. Mating behaviour in crustaceans could potentially be severely affected by anthropogenic noise, given that noise has been demonstrated to impact some crustacean's ability to detect and respond to chemical, visual, and acoustic cues, all of which are vital in courtship rituals. To explore if noise has an impact on crustacean mating, we tested the responses of male green shore crabs (Carcinus maenas) from the southwest UK coast by exposing them to ship noise recordings while simultaneously presenting them with a dummy-female soaked in the female-sex pheromone uridine diphosphate (UDP) in an experimental tank setup (recording treatment: n = 15, control treatment: n = 15). We found a significant, negative effect of noise on the occurrence of mating behaviour compared to no noise conditions, though no significant effect of noise on the time it took for a crab to respond to the pheromone. Such effects suggest reproductive impairment due to anthropogenic noise, which could potentially contribute to decreased crustacean populations and subsequent ecological and economic repercussions. Given the findings of our preliminary study, more research should be undertaken that includes larger sample sizes, double blind setups, and controlled laboratory trials in order to more fully extrapolate the potential impact of noise on mating in the natural environment.

Adapting to climate with limited genetic diversity: Nucleotide, DNA methylation and microbiome variation among populations of the social spider Stegodyphus dumicola.

Understanding the role of genetic and nongenetic variants in modulating phenotypes is central to our knowledge of adaptive responses to local conditions and environmental change, particularly in species with such low population genetic diversity that it is likely to limit their evolutionary potential. A first step towards uncovering the molecular mechanisms underlying population-specific responses to the environment is to carry out environmental association studies. We associated climatic variation with genetic, epigenetic and microbiome variation in populations of a social spider with extremely low standing genetic diversity. We identified genetic variants that are associated strongly with environmental variation, particularly with average temperature, a pattern consistent with local adaptation. Variation in DNA methylation in many genes was strongly correlated with a wide set of climate parameters, thereby revealing a different pattern of associations than that of genetic variants, which show strong correlations to a more restricted range of climate parameters. DNA methylation levels were largely independent of cis-genetic variation and of overall genetic population structure, suggesting that DNA methylation can work as an independent mechanism. Microbiome composition also correlated with environmental variation, but most strong associations were with precipitation-related climatic factors. Our results suggest a role for both genetic and nongenetic mechanisms in shaping phenotypic responses to local environments.

Estimating cetacean population trends from static acoustic monitoring data using Paired Year Ratio Assessment (PYRA).

The cetacean conservationist is often faced with evaluating population trends from abundance data that are either sparse or recorded at different times in different years. The presence of diel or seasonal patterns in the data together with unplanned gaps is often problematic. Such data are typical of those obtained from static acoustic monitoring. We present a simple and transparent non-parametric trend evaluation method, 'Paired Year Ratio Assessment (PYRA)' that uses only whole days of data wherever they are present in each of successive pairs of periods of 365 days. We provide a quantitative comparison of the performance of PYRA with traditional generalised additive models (GAMS) and nonparametric randomisation tests that require a greater level of skill and experience for both application and interpretation. We conclude that PYRA is a powerful tool, particularly in the context of identifying population trends which is often the main aim of conservation-targeted acoustic monitoring.

Telomere length is highly heritable and independent of growth rate manipulated by temperature in field crickets.

Many organisms are capable of growing faster than they do. Restrained growth rate has functionally been explained by negative effects on lifespan of accelerated growth. However, the underlying mechanisms remain elusive. Telomere attrition has been proposed as a causal agent and has been mostly studied in endothermic vertebrates. We established that telomeres exist as chromosomal-ends in a model insect, the field cricket Gryllus campestris, using terminal restriction fragment and Bal 31 methods. Telomeres comprised TTAGGn repeats of 38 kb on average, more than four times longer than the telomeres of human infants. Bal 31 assays confirmed that telomeric repeats were located at the chromosome-ends. We tested whether rapid growth between day 1, day 65, day 85, and day 125 is achieved at the expense of telomere length by comparing nymphs reared at 23°C with their siblings reared at 28°C, which grew three times faster in the initial 65 days. Surprisingly, neither temperature treatment nor age affected average telomere length. Concomitantly, the broad sense heritability of telomere length was remarkably high at ~100%. Despite high heritability, the evolvability (a mean-standardized measure of genetic variance) was low relative to that of body mass. We discuss our findings in the context of telomere evolution. Some important features of vertebrate telomere biology are evident in an insect species dating back to the Triassic. The apparent lack of an effect of growth rate on telomere length is puzzling, suggesting strong telomere length maintenance during the growth phase. Whether such maintenance of telomere length is adaptive remains elusive and requires further study investigating the links with fitness in the wild.

Evidence for genetic isolation and local adaptation in the field cricket Gryllus campestris.

Understanding how species can thrive in a range of environments is a central challenge for evolutionary ecology. There is strong evidence for local adaptation along large-scale ecological clines in insects. However, potential adaptation among neighbouring populations differing in their environment has been studied much less. We used RAD sequencing to quantify genetic divergence and clustering of ten populations of the field cricket Gryllus campestris in the Cantabrian Mountains of northern Spain, and an outgroup on the inland plain. Our populations were chosen to represent replicate high and low altitude habitats. We identified genetic clusters that include both high and low altitude populations indicating that the two habitat types do not hold ancestrally distinct lineages. Using common-garden rearing experiments to remove environmental effects, we found evidence for differences between high and low altitude populations in physiological and life-history traits. As predicted by the local adaptation hypothesis, crickets with parents from cooler (high altitude) populations recovered from periods of extreme cooling more rapidly than those with parents from warmer (low altitude) populations. Growth rates also differed between offspring from high and low altitude populations. However, contrary to our prediction that crickets from high altitudes would grow faster, the most striking difference was that at high temperatures, growth was fastest in individuals from low altitudes. Our findings reveal that populations a few tens of kilometres apart have independently evolved adaptations to their environment. This suggests that local adaptation in a range of traits may be commonplace even in mobile invertebrates at scales of a small fraction of species' distributions.

Ship noise inhibits colour change, camouflage, and anti-predator behaviour in shore crabs.

Ship noise is a prominent source of underwater sound pollution. Carter et al. demonstrate that ship noise has multiple negative effects on animal traits that do not primarily rely on acoustics. In shore crabs, color change to improve camouflage and predator escape responses are adversely affected by ship noise but not by equally loud ambient noise.

Larval Host Preference and Suitability for the Sawfly Mesoneura rufonota among Five Cinnamomun Tree Species.

The camphor sawfly, Mesoneura rufonota Rohwer, is an economically important leaf-chewing pest of the genus Cinnamomum Trew. However, little is known about the range of species that it can infest within this genus or whether larvae show preferences for particular species. We conducted preference and performance experiments under laboratory conditions to assess larval host preference and suitability as a host plant of five congeneric trees species: C. camphora (Linn) Presl, C. bodinieri Levl., C. burmanni (Nees et T. Nees) Blume, C. pauciflorum Nees, and C. micranthum (Hay.) Hay. In no-choice, two-choice and multiple-choice feeding trials, significantly higher feeding rates were found on C. camphora, followed by C. bodinieri, compared to those on the other three tree species. In two-choice behavior trials, larvae moved to occupy leaves of C. camphora faster and more frequently, followed by C. bodinieri, than when offered the other three tree species. In no-choice fitness trials, the survival of larval and pupal stage was the highest, the developmental duration of larval and pupal stage was the shortest, the pupal body weight was the heaviest and adults lived the longest on C. camphora followed by C. bodinieri, while M. rufonota did not complete development on C. burmanni, C. pauciflorum or C. micranthum. Our results demonstrate that larvae consistently prefer and perform better on C. camphora leaves, that they can utilize C. bodinieri, but it is less preferred, and that C. burmanni, C. pauciflorum, and C. micranthum appear to be unsuitable as a host for M. rufonota.

Background matching and disruptive coloration as habitat-specific strategies for camouflage.

Camouflage is a key defence across taxa and frequently critical to survival. A common strategy is background matching, resembling the colour and pattern of the environment. This approach, however, may be ineffective in complex habitats where matching one patch may lead to increased visibility in other patches. In contrast, disruptive coloration, which disguises body outlines, may be effective against complex backgrounds. These ideas have rarely been tested and previous work focuses on artificial systems. Here, we test the camouflage strategies of the shore crab (Carcinus maenas) in two habitats, being a species that is highly variable, capable of plastic changes in appearance, and lives in multiple environments. Using predator (bird and fish) vision modelling and image analysis, we quantified background matching and disruption in crabs from rock pools and mudflats, predicting that disruption would dominate in visually complex rock pools but background matching in more uniform mudflats. As expected, rock pool individuals had significantly higher edge disruption than mudflat crabs, whereas mudflat crabs more closely matched the substrate than rock pool crabs for colour, luminance, and pattern. Our study demonstrates facultative expression of camouflage strategies dependent on the visual environment, with implications for the evolution and interrelatedness of defensive strategies.

Slower senescence in a wild insect population in years with a more female-biased sex ratio.

Life-history theories of senescence are based on the existence of a trade-off in resource allocation between body maintenance and reproduction. This putative trade-off means that environmental and demographic factors affecting the costs of reproduction should be associated with changes in patterns of senescence. In many species, competition among males is a major component of male reproductive investment, and hence variation in the sex ratio is expected to affect rates of senescence. We test this prediction using nine years of demographic and behavioural data from a wild population of the annual field cricket Gryllus campestris. Over these generations, the sex ratio at adulthood varied substantially, from years with an equal number of each sex to years with twice as many females as males. Consistent with the predictions of theory, we found that in years with a greater proportion of females, both sexes experienced a slower increase in mortality rate with age. Additionally, phenotypic senescence in males was slower in years when there were more females. Sex ratio did not affect the baseline mortality rate in males, but females suffered higher age-independent mortality rates when males were in short supply.

Comparing individual and population measures of senescence across 10 years in a wild insect population.

Declines in survival and performance with advancing age (senescence) have been widely documented in natural populations, but whether patterns of senescence across traits reflect a common underlying process of biological ageing remains unclear. Senescence is typically characterized via assessments of the rate of change in mortality with age (actuarial senescence) or the rate of change in phenotypic performance with age (phenotypic senescence). Although both phenomena are considered indicative of underlying declines in somatic integrity, whether actuarial and phenotypic senescence rates are actually correlated has yet to be established. Here we present evidence of both actuarial and phenotypic senescence from a decade-long longitudinal field study of wild insects. By tagging every individual and using continuous video monitoring with a network of up to 140 video cameras, we were able to record survival and behavioral data on an entire adult population of field crickets. This reveals that both actuarial and phenotypic senescence vary substantially across 10 annual generations. This variation allows us to identify a strong correlation between actuarial and phenotypic measures of senescence. Our study demonstrates age-related phenotypic declines reflected in population level mortality rates and reveals that observations of senescence in a single year may not be representative of a general pattern.

Testing the effect of early-life reproductive effort on age-related decline in a wild insect.

The disposable soma theory of ageing predicts that when organisms invest in reproduction they do so by reducing their investment in body maintenance, inducing a trade-off between reproduction and survival. Experiments on invertebrates in the lab provide support for the theory by demonstrating the predicted responses to manipulation of reproductive effort or lifespan. However, experimental studies in birds and evidence from observational (nonmanipulative) studies in nature do not consistently reveal trade-offs. Most species studied previously in the wild are mammals and birds that reproduce over multiple discrete seasons. This contrasts with temperate invertebrates, which typically have annual generations and reproduce over a single season. We expand the taxonomic range of senescence study systems to include life histories typical of most temperate invertebrates. We monitored reproductive effort, ageing, and survival in a natural field cricket population over ten years to test the prediction that individuals investing more in early-reproduction senesce faster and die younger. We found no evidence of a trade-off between early-life reproductive effort and survival, and only weak evidence for a trade-off with phenotypic senescence. We discuss the possibility that organisms with multiple discrete breeding seasons may have greater opportunities to express trade-offs between reproduction and senescence.

Reproductive isolation in the acoustically divergent groups of tettigoniid, Mecopoda elongata.

Sympatric divergent populations of the same species provide an opportunity to study the evolution and maintenance of reproductive isolation. Male mating calls are important in sexual selection in acoustically communicating species, and they also have the potential to maintain isolation among species or incipient species. We studied divergent south Indian populations of the bush cricket Mecopoda elongata which are extremely difficult to distinguish morphologically, but which exhibit striking divergence in male acoustic signals. We performed phonotactic experiments investigating the relative preference of females of the "Chirper" song type for calls of all 5 of the song types found in the region (in varying degrees of sympatry). We found that Chirper females preferred their own song type and were completely unresponsive to three trilling song types. Chirper females were occasionally attracted to the call type "Double Chirper" (the call most similar to their own type), suggesting call preference alone cannot provide a complete isolating mechanism. To investigate the basis of call preference we investigated the response of chirper females to variation in chirp rate. Chirper females responded most frequently to a mean chirp rate characteristic of their own song type rather than a higher chirp rate which would be more characteristic of the Double-Chirper song type. This suggests females drive stabilising selection on male chirp rate, which may contribute to the maintenance of isolation. Finally, a no-choice mating experiment using Chirper females and Chirper and Double Chirper males revealed a significant preference of Chirper females to mate with their own song type, even without a requirement for phonotaxis. Overall, the strong specificity of Chirper females for their 'own' song type provides evidence for behavioural isolation among divergent sympatric Mecopoda song types being maintained by female preference for both male song type and subsequent mating probability driven by other cues.

Analysing animal social network dynamics: the potential of stochastic actor-oriented models.

Animals are embedded in dynamically changing networks of relationships with conspecifics. These dynamic networks are fundamental aspects of their environment, creating selection on behaviours and other traits. However, most social network-based approaches in ecology are constrained to considering networks as static, despite several calls for such analyses to become more dynamic. There are a number of statistical analyses developed in the social sciences that are increasingly being applied to animal networks, of which stochastic actor-oriented models (SAOMs) are a principal example. SAOMs are a class of individual-based models designed to model transitions in networks between discrete time points, as influenced by network structure and covariates. It is not clear, however, how useful such techniques are to ecologists, and whether they are suited to animal social networks. We review the recent applications of SAOMs to animal networks, outlining findings and assessing the strengths and weaknesses of SAOMs when applied to animal rather than human networks. We go on to highlight the types of ecological and evolutionary processes that SAOMs can be used to study. SAOMs can include effects and covariates for individuals, dyads and populations, which can be constant or variable. This allows for the examination of a wide range of questions of interest to ecologists. However, high-resolution data are required, meaning SAOMs will not be useable in all study systems. It remains unclear how robust SAOMs are to missing data and uncertainty around social relationships. Ultimately, we encourage the careful application of SAOMs in appropriate systems, with dynamic network analyses likely to prove highly informative. Researchers can then extend the basic method to tackle a range of existing questions in ecology and explore novel lines of questioning.

Wild cricket social networks show stability across generations.

BACKGROUND: A central part of an animal's environment is its interactions with conspecifics. There has been growing interest in the potential to capture these interactions in the form of a social network. Such networks can then be used to examine how relationships among individuals affect ecological and evolutionary processes. However, in the context of selection and evolution, the utility of this approach relies on social network structures persisting across generations. This is an assumption that has been difficult to test because networks spanning multiple generations have not been available. We constructed social networks for six annual generations over a period of eight years for a wild population of the cricket Gryllus campestris. RESULTS: Through the use of exponential random graph models (ERGMs), we found that the networks in any given year were able to predict the structure of networks in other years for some network characteristics. The capacity of a network model of any given year to predict the networks of other years did not depend on how far apart those other years were in time. Instead, the capacity of a network model to predict the structure of a network in another year depended on the similarity in population size between those years. CONCLUSIONS: Our results indicate that cricket social network structure resists the turnover of individuals and is stable across generations. This would allow evolutionary processes that rely on network structure to take place. The influence of network size may indicate that scaling up findings on social behaviour from small populations to larger ones will be difficult. Our study also illustrates the utility of ERGMs for comparing networks, a task for which an effective approach has been elusive.

Comparing pre- and post-copulatory mate competition using social network analysis in wild crickets.

Sexual selection results from variation in success at multiple stages in the mating process, including competition before and after mating. The relationship between these forms of competition, such as whether they trade-off or reinforce one another, influences the role of sexual selection in evolution. However, the relationship between these 2 forms of competition is rarely quantified in the wild. We used video cameras to observe competition among male field crickets and their matings in the wild. We characterized pre- and post-copulatory competition as 2 networks of competing individuals. Social network analysis then allowed us to determine 1) the effectiveness of precopulatory competition for avoiding postcopulatory competition, 2) the potential for divergent mating strategies, and 3) whether increased postcopulatory competition reduces the apparent reproductive benefits of male promiscuity. We found 1) limited effectiveness of precopulatory competition for avoiding postcopulatory competition; 2) males do not specifically engage in only 1 type of competition; and 3) promiscuous individuals tend to mate with each other, which will tend to reduce variance in reproductive success in the population and highlights the trade-off inherent in mate guarding. Our results provide novel insights into the works of sexual competition in the wild. Furthermore, our study demonstrates the utility of using network analyses to study competitive interactions, even in species lacking obvious social structure.

Dynamics of among-individual behavioral variation over adult lifespan in a wild insect.

Investigating patterns of among and within-individual trait variation in populations is essential to understanding how selection shapes phenotypes. Behavior is often the most flexible aspect of the phenotype, and to understand how it is affected by selection, we need to examine how consistent individuals are. However, it is not well understood whether among-individual differences tend to remain consistent over lifetimes, or whether the behavior of individuals relative to one another varies over time. We examined the dynamics of 4 behavioral traits (tendency to leave a refuge, shyness, activity, and exploration) in a wild population of field crickets (Gryllus campestris). We tagged individuals and then temporarily removed them from their natural environment and tested them under laboratory conditions. All 4 traits showed among-individual variance in mean levels of expression across the adult lifespan, but no significant differences in how rapidly expression changed with age. For all traits, among-individual variance increased as individuals got older. Our findings reveal seldom examined changes in variance components over the adult lifetime of wild individuals. Such changes will have important implications for the relationship between behavioral traits, life-histories, and fitness and the consequences of selection on wild individuals.

Behaviour in captivity predicts some aspects of natural behaviour, but not others, in a wild cricket population.

Examining the relevance of 'animal personality' involves linking consistent among- and within-individual behavioural variation to fitness in the wild. Studies aiming to do this typically assay personality in captivity and rely on the assumption that measures of traits in the laboratory reflect their expression in nature. We examined this rarely tested assumption by comparing laboratory and field measurements of the behaviour of wild field crickets (Gryllus campestris) by continuously monitoring individual behaviour in nature, and repeatedly capturing the same individuals and measuring their behaviour in captivity. We focused on three traits that are frequently examined in personality studies: shyness, activity and exploration. All of them showed repeatability in the laboratory. Laboratory activity and exploration predicted the expression of their equivalent behaviours in the wild, but shyness did not. Traits in the wild were predictably influenced by environmental factors such as temperature and sunlight, but only activity showed appreciable within-individual repeatability. This suggests that some behaviours typically studied as personality traits can be accurately assayed in captivity, but the expression of others may be highly context-specific. Our results highlight the importance of validating the relevance of laboratory behavioural assays to analogous traits measured in the wild.

Evolutionary rates for multivariate traits: the role of selection and genetic variation.

A fundamental question in evolutionary biology is the relative importance of selection and genetic architecture in determining evolutionary rates. Adaptive evolution can be described by the multivariate breeders' equation (Δz(-)=Gß), which predicts evolutionary change for a suite of phenotypic traits (Δz(-)) as a product of directional selection acting on them (ß) and the genetic variance-covariance matrix for those traits (G ). Despite being empirically challenging to estimate, there are enough published estimates of G and ß to allow for synthesis of general patterns across species. We use published estimates to test the hypotheses that there are systematic differences in the rate of evolution among trait types, and that these differences are, in part, due to genetic architecture. We find some evidence that sexually selected traits exhibit faster rates of evolution compared with life-history or morphological traits. This difference does not appear to be related to stronger selection on sexually selected traits. Using numerous proposed approaches to quantifying the shape, size and structure of G, we examine how these parameters relate to one another, and how they vary among taxonomic and trait groupings. Despite considerable variation, they do not explain the observed differences in evolutionary rates.