Transgenerational plasticity (TGP) has largely focused on how parental exposure to ecological conditions shapes the phenotypes of future generations. However, organisms acquire information about their ecological environment via social learning, which can also shape TGP in profound ways. We demonstrate that non-parents alter how parents detect and respond to environmental cues in ways that spillover to affect offspring, non-parents influence offspring even without direct physical interactions, and parental cues received by offspring can alter the phenotypes of other juveniles. Because parents can draw on the experiences of a network of non-parents, these socially acquired cues may increase parents’ ability to accurately detect environmental shifts and may explain why TGP is surprisingly ubiquitous despite theory predicting that it should be relatively rare.

Plastic pollution threatens almost every ecosystem in the world. Critically, many animals consume plastic, in part because plastic particles often look or smell like food. Plastic ingestion is thus an evolutionary trap, a phenomenon that occurs when cues are decoupled from their previously associated high fitness outcomes. Theory predicts that dominance hierarchies could dictate individual responses to evolutionary traps across social environments, but the social dimension of evolutionary trap responses has rarely been investigated. We tested how variation in group size influences the formation of dominance relationships and, in turn, how these dominance relationships drive differences in foraging behavior in Western mosquitofish (Gambusia affinis). This included foraging for a variety of familiar and novel food-like items, including microplastics. Overall, dominant individuals were often the first to sample food and had higher bite rates than subordinates, including when foraging for microplastics. Importantly, how dominance affected foraging behavior depended on group size and on whether groups were presented with familiar or novel foods. Furthermore, individuals were consistent in their foraging behavior across trials with different group sizes, indicating the formation of stable social roles. These results suggest that predicting the ecological and evolutionary consequences of evolutionary traps will require an understanding of how social structures influence trap susceptibility.

Climate change will affect both the mean and the variability in environmental conditions and may have major negative impacts on population densities in the future. For annual plants that already live in an extreme environment like the Sonoran Desert, keeping a fraction of their seeds dormant underground (for possibly years at a time) is critical to survive. Here, we consider how this form of bet hedging (i.e., delayed germination) for 10 Sonoran Desert annuals mediates responses to precipitation shifts. We use a demographic model parameterized with long-term field and precipitation data to explore how forecasted changes in precipitation impact annual plant species' population densities. We then examine how instantaneous evolution of optimal germination fractions in the shifted precipitation regimes bolsters population densities. Our results indicate that overall less rainfall and, to a lesser extent, increased variance in rainfall drive population levels down. Instantaneous evolution of optimal germination fractions in new regimes benefited species' populations only marginally, and only for small to moderate shifts in precipitation. Thus, even rapid evolution is unlikely to save populations experiencing larger shifts in precipitation. Finally, we predict that specialists that can capitalize on wet-year bonanzas or are water use efficient will be the most resilient to precipitation shifts as long as their seed survivorships are sufficiently high.

The tendency of wildlife to associate with humans spatially and temporally, i.e., to overlap with humans, is a key factor mediating human-wildlife coexistence in cities. While direct human influence (e.g., food subsidies) is recognized as a mechanism of wildlife overlap with humans, the role of broader urban ecosystem characteristics (e.g., habitat availability, human sociodemographics) in shaping this behavior is a burgeoning area of research. We analyzed the relationship between environmental and societal characteristics and overlap with humans in coyotes, a species of management concern due to emerging human-coyote conflicts in cities across North America. Using high-resolution GPS data from coyotes in the Chicago metropolitan area, we estimated overlap by quantifying coyotes’ spatial selection for human population density during periods of temporal overlap, i.e., periods when coyotes and humans were active. On average, coyotes exhibited high overlap, evidenced by their selection for areas of high human population density during periods of temporal overlap, but with substantial individual variation in this selection. Importantly, the mean level of overlap depended on environmental characteristics. Selection for human population density was negatively associated with the level of impervious surface cover and positively associated with the presence of land uses containing natural habitat features (e.g., parks, golf courses, vacant lots). Coyote selection for human population density was not significantly associated with median income or the proportion of white residents. Our results suggest that natural habitat features in densely populated areas may be hotspots for human-coyote interactions. Efforts to manage human-coyote interactions should focus on these areas and identify individual traits that heighten coyote overlap with humans.

Theory predicts that high population density leads to more strongly connected spatial and social networks, but how local density drives individuals' positions within their networks is unclear. This gap reduces our ability to understand and predict density-dependent processes. Here we show that density drives greater network connectedness at the scale of individuals within wild animal populations. Across 36 datasets of spatial and social behaviour in >58,000 individual animals, spanning 30 species of fish, reptiles, birds, mammals and insects, 80% of systems exhibit strong positive relationships between local density and network centrality. However, >80% of relationships are nonlinear and 75% are shallower at higher values, indicating saturating trends that probably emerge as a result of demographic and behavioural processes that counteract density's effects. These are stronger and less saturating in spatial compared with social networks, as individuals become disproportionately spatially connected rather than socially connected at higher densities. Consequently, ecological processes that depend on spatial connections are probably more density dependent than those involving social interactions. These findings suggest fundamental scaling rules governing animal social dynamics, which could help to predict network structures in novel systems.

Host personality can markedly affect parasite transmission. Especially for parasites with indirect transmission through the environment, the effects of consistent among-individual differences in behavior may have both direct and indirect components. For example, personality may mediate both how hosts respond to infected individuals and the likelihood that hosts indirectly interact with infected conspecifics (e.g., by visiting patches infected hosts have previously contaminated). Integrating parasites, personality, and these different kinds of interaction networks constitutes a key step toward understanding transmission in natural systems. We evaluated these elements using a 5-year field study of a wild population of sleepy lizards, Tiliqua rugosa, and their tick parasites, which transmit among lizards through lizards' shared use of refuges. Using Bayesian models, we evaluated (1) predictors of lizard infestation probability and intensity (i.e., average tick count when infested) and (2) relationships among the predictors. We used the latter set of models to assess indirect relationships between the predictors and the infestation metrics. As predictors, we used lizards' infestation "risk" (derived from a time-lagged refuge sharing transmission network), traits (sex, mass, and the personality axes aggression and boldness), space use (number of unique refuges used and home range overlap with other lizards), and measures of synchronous social interactions (i.e., edge weight and degree). We found both indirect and direct connections between our predictors and tick infestation. For example, boldness was positively directly associated with infection intensity and indirectly positively associated with both infestation probability and intensity via intermediary connections with social network interaction and risk. Using more unique refuges, on the other hand, was indirectly negatively associated with infestation probability (via reduced risk), but directly positively associated with infestation probability, indicating a potential trade-off in the anti-parasite benefits of using more refuges. Our results emphasize that (1) multiple aspects of host behavior may associate with parasite infection, (2) these components may proceed through both direct and indirect pathways, and (3) multiple pathways should be considered together because the pathways may have compounding or counteracting effects.

Humans may play a key role in providing small prey mammals spatial and temporal refuge from predators, but few studies have captured the heterogeneity of these effects across space and time. Global COVID-19 lockdown restrictions offered a unique opportunity to investigate how a sudden change in human presence in a semi-urban park impacted wildlife. Here, we quantify how changes in the spatial distributions of humans and natural predators influenced the landscape of fear for the California ground squirrel (Otospermophilus beecheyi) in a COVID-19 pandemic (2020) and non-COVID (2019) year. We used a structural equation modeling approach to explore the direct and indirect effects of human presence, predator presence, and habitat features on foraging that reflected fear responses (e.g., giving-up densities [GUDs], number of foragers, and average food intake rate while at food patches). In 2019, humans and dogs had moderate effects on GUDs; squirrels were less fearful (lower GUDs) in areas frequently visited by humans and dogs, but the effects of raptors were weak. In contrast, in 2020, the effects of humans and dogs on GUDs were weak; squirrels were more fearful of high raptor activity, open sky, and ground cover. In both years, squirrels farthest from refuge were the most risk-averse. Overall, our analyses revealed an increase in perceived risk from natural predators in 2020 associated with a change in the concentration of human presence. Thus, risk-sensitive foraging was dynamic across space and time, depending on a complex interplay among human and dog activity, natural predators, and microhabitat features. Our findings elucidate the myriad ways humans directly and indirectly influence animal perception of safety and danger.

Host behavior and parasite infection reciprocally interact, but this dynamic is rarely studied experimentally in the field with multiple behaviors. We investigated the interplay between parasitism and host behavior via an in situ experimental tick infestation of a wild population of sleepy lizards, Tiliqua rugosa. Using Bayesian models, we assessed the relationship between experimental infestation and lizard aggression and boldness before and after infestation. First, we tested whether lizard aggression and boldness prior to infestation predicted the probability of tick attachment in the infestation experiment. Second, we evaluated whether experimental infestation affected subsequent lizard aggression and boldness. We found that aggression and boldness related interactively with infestation: for unaggressive lizards, higher boldness was associated with reduced experimental infestation success, but the opposite occurred for aggressive individuals. Second, increased tick infestation did not affect post-infestation aggression, but tended to increase boldness. Taken together, these results highlight the potential for feedbacks between parasites and multi-dimensional host behaviors. Although behavior is known to affect parasite loads, and parasites, in turn, can affect host behavior, it is unclear how personality interacts with parasite infestation. We experimentally infected a wild lizard population with ticks to study how lizard personality interacted with parasite loads in complex feedback loops. We found that personality affected parasite loads and that parasites, in turn, affected hosts with different personality types in distinct ways.

Anti-predator behaviours in response to predator cues can be innate, or they can be learned through prior experience and remembered over time. The duration and strength of continued anti-predator behaviour after predator cues are no longer present, and the potential for an enhanced response when re-exposed to predator cues later is less known but could account for the observed variation in anti-predator responses. We measured the carryover effects of past predation exposure and the potential for anti-predator learning and memory in the marine snail Nucella canaliculata from six populations distributed over 1000 km of coastline. We exposed lab-reared snails to cues associated with a common crab predator or seawater control in two serial experiments separated by over seven months. Responses were population- and sex-dependent, with some populations retaining anti-predator behaviours while others showed a capacity for learning and memory. Male snails showed a strong carryover of risk aversion, while females were able to return to normal feeding rates and grow more quickly. These behavioural differences culminated in strong impacts on feeding and growth rates, demonstrating that this variation has implications for the strength of trait-mediated indirect interactions, which can impact entire ecosystems.

Animal gut microbiomes can be very diverse, and enteric bacteria can profoundly affect the physiology of their host. The gut microbiome can be related to host health and digestion, which ultimately contribute to host body condition. However, we have a limited understanding of the co-occurrence patterns of gut bacteria in their host, and how co-occurrence and bacterial diversity change over time. This notion is especially important to animals living in groups as bacteria can transmit through social interactions. We investigated the co-occurrence patterns of gut bacteria in a lizard host. We repeatedly collected cloacal swabs from 87 sleepy lizards (Tiliqua rugosa) from two different study sites over their activity season. We determined the richness and prevalence of 82 enteric bacterial strains and used a probabilistic model to investigate their co-occurrence. At both study sites, richness and prevalence generally increased over time. We suggest that the lizards acquire strains throughout their activity season by moving through the landscape and inspecting conspecific scats. Lizards continuously tongue-flick while moving, and thereby ingest bacteria when they move through areas where other animals defaecated. Temperature, rainfall and diet change seasonally, influencing lizard activity, and may influence the observed increase in enterobacterial richness and prevalence. Further, albeit with some exceptions, most strain pairs did not occur significantly more often or less often than expected by chance. This finding shows a lack of structured co-occurrence, which may imply that most bacterial strains did not facilitate or inhibit each other. The absence of a co-occurrence pattern could also be driven by random encounters of bacteria shed by other lizards within the habitat. Our results suggest that behaviour (movement patterns, tongue-flicking), activity patterns and environmental factors collectively drive the temporal pattern of the gut bacterial community in sleepy lizards and potentially other wild reptiles.

While numerous studies have examined either initial prey responses to an approaching predator (flight initiation distance, FID), or subsequent hiding behavior (e.g. latency to resume activity), to our knowledge, no previous studies have repeatedly tested multiple individuals in nature, to quantify whether both FIDs and latencies to resume activity are repeatable, and whether these two stages of the antipredator response are positively correlated. This correlation is ecologically important in that opportunity costs of predator avoidance are particularly large if the same individuals tend to both escape more readily and hide longer. Here, we examined California ground squirrels’ (Otospermophilus beecheyi) responses to human approach, and provided the first example showing that, as predicted: FIDs, latencies to resume activity, and other aspects of prey responses post-FID were repeatable and positively correlated. Interestingly, we also found that across a gradient of human activity, squirrels in areas with higher human activity were generally bolder.

Recent evidence suggests that individuals differ in foraging tactics and this variation is often linked to an individual's behavioural type (BT). Yet, while foraging typically comprises a series of search and handling steps, empirical investigations have rarely considered BT-dependent effects across multiple stages of the foraging process, particularly in natural settings. In our long-term sleepy lizard (Tiliqua rugosa) study system, individuals exhibit behavioural consistency in boldness (measured as an individual's willingness to approach a novel food item in the presence of a threat) and aggressiveness (measured as an individual's response to an 'attack' by a conspecific dummy). These BTs are only weakly correlated and have previously been shown to have interactive effects on lizard space use and movement, suggesting that they could also affect lizard foraging performance, particularly in their search behaviour for food. To investigate how lizards' BTs affect their foraging process in the wild, we supplemented food in 123 patches across a 120-ha study site with three food abundance treatments (high, low and no-food controls). Patches were replenished twice a week over the species' entire spring activity season and feeding behaviours were quantified with camera traps at these patches. We tracked lizards using GPS to determine their home range (HR) size and repeatedly assayed their aggressiveness and boldness in designated assays. We hypothesised that bolder lizards would be more efficient foragers while aggressive ones would be less attentive to the quality of foraging patches. We found an interactive BT effect on overall foraging performance. Individuals that were both bold and aggressive ate the highest number of food items from the foraging array. Further dissection of the foraging process showed that aggressive lizards in general ate the fewest food items in part because they visited foraging patches less regularly, and because they discriminated less between high and low-quality patches when revisiting them. Bolder lizards, in contrast, ate more tomatoes because they visited foraging patches more regularly, and ate a higher proportion of the available tomatoes at patches during visits. Our study demonstrates that BTs can interact to affect different search and handling components of the foraging process, leading to within-population variation in foraging success. Given that individual differences in foraging and movement will influence social and ecological interactions, our results highlight the potential role of BT's in shaping individual fitness strategies and population dynamics.

Survival links individual-level responses to population, community, and ecosystem dynamics. Thus, understanding species’ survival responses to environmental change in urban areas is critical for gaining insights into the ecology and management of wildlife in these rapidly expanding environments. Despite existing research on the broadscale effects of urbanization on wildlife survival, the impacts of heterogeneity within urban areas remain largely unexplored. We identified environmental and societal characteristics evidenced to influence the distribution of resources and risks important for urban wildlife ecology. Using Cox proportional hazards models, we examined the association between these characteristics and the survival of coyotes (Canis latrans) living in the Chicago metropolitan area. We found a significant interaction between income and human population density, such that survival was negatively associated with income in densely populated areas. Notably, no environmental characteristics were significantly associated with survival. This study adds to the growing literature demonstrating the significance of ecological heterogeneity in wildlife responses to urbanization and highlights society’s influence on wildlife survival in cities.

Outdoor access may provide mental stimulation and enrichment for companion cats (Felis catus); however, it also raises concerns regarding their safety and predation of vulnerable local wildlife. Understanding factors that shape cats' outdoor access and behavioral outcomes is therefore crucial for identifying nuanced solutions to the human-cat-wildlife conflict. In this exploratory study, we aimed to 1) explore owner attitudes towards providing or reducing outdoor access, 2) identify general predictors of free-roaming outdoor access, and 3) identify predictors of hunting behavior among cats that are allowed outside. Using a quantitative questionnaire (n=75 questions), we collected information on cat owners' demographics and attitudes, their cat's characteristics and behaviors in the home, home environment, outdoor lifestyle, and hunting history. Of 631 cat owners living near Davis, California, USA, recruited through convenience and snowball sampling, 36.7 % indicated they were keeping their cat indoor-only (n=232), 34.1 % providing supervised or controlled outdoor access (n=215), and 29.2 % providing uncontrolled, or free-roaming outdoor access (n=184). More than half of outdoor cat owners reported they were likely or somewhat likely to reduce outdoor access if they lived near busier streets (59 %, n=235), if more dangerous animals were around (59 %, n=235), if their cat stopped coming home consistently (56 %, n=223), or if they knew their cat was negatively impacting wildlife (53 %, n=211). We found that cats had higher odds of freely roaming if they were 7-10 years old compared to <1 year old (p=0.030), obtained from the street rather than a shelter (p=0.007), living within 500 ft of traffic (p=0.002), living with a dog (p=0.014) or a child (p=0.005), or if they showed aggressive behaviors in the home (p=0.008). Cats had lower odds of freely roaming if they had scratching areas in the home (p=0.017), or were living with one (p=0.020) or two (p=0.017) other cats. Among those allowed outside, cats had higher odds of having hunted prey in the last year if they were allowed to freely roam without supervision (p=0.016), or were allowed outdoors between 6:00-11:59 am (p=0.014). These results generate directions for future research in the field of companion animal management, and emphasize the role of cat owners in the intersection of animal welfare and wildlife conservation.

High population density should drive individuals to more frequently share space and interact, producing better-connected spatial and social networks. Despite this widely-held assumption, it remains unconfirmed how local density generally drives individuals' positions within wild animal networks. We analysed 34 datasets of simultaneous spatial and social behaviour in >55,000 individual animals, spanning 28 species of fish, reptiles, birds, mammals, and insects. >80% of systems exhibited strongly positive relationships between local density and network centrality, providing broad empirical evidence that local density increases connectedness at the individual level. However, >75% of density-connectedness relationships were nonlinear, and density's importance declined at higher values in >70% of systems, signifying saturating effects. Density's effect was much stronger and less saturating for spatial than social networks, suggesting population density drives individuals to become disproportionately spatially connected rather than socially. These findings reveal fundamental trends underlying societal structuring, with widespread behavioural, ecological, and evolutionary implications. ### Competing Interest Statement The authors have declared no competing interest.

Fire regimes are changing dramatically worldwide due to climate change, habitat conversion, and the suppression of Indigenous landscape management. Although there has been extensive work on plant responses to fire, including their adaptations to withstand fire and long-term effects of fire on plant communities, less is known about animal responses to fire. Ecologists lack a conceptual framework for understanding behavioural responses to fire, which can hinder wildlife conservation and management. Here, we integrate cue-response sensory ecology and predator-prey theory to predict and explain variation in if, when and how animals react to approaching fire. Inspired by the literature on prey responses to predation risk, this framework considers both fire-naïve and fire-adapted animals and follows three key steps: vigilance, cue detection and response. We draw from theory on vigilance tradeoffs, signal detection, speed-accuracy tradeoffs, fear generalization, neophobia and adaptive dispersal. We discuss how evolutionary history with fire, but also other selective pressures, such as predation risk, should influence animal behavioural responses to fire. We conclude by providing guidance for empiricists and outlining potential conservation applications.

While a large body of research has focused on the physiological effects of multiple environmental stressors, how behavioural and life-history plasticity mediate multiple-stressor effects remains underexplored. Behavioural plasticity can not only drive organism-level responses to stressors directly but can also mediate physiological responses. Here, we provide a conceptual framework incorporating four fundamental trade-offs that explicitly link animal behaviour to life-history-based pathways for energy allocation, shaping the impact of multiple stressors on fitness. We first address how small-scale behavioural changes can either mediate or drive conflicts between the effects of multiple stressors and alternative physiological responses. We then discuss how animal behaviour gives rise to three additional understudied and interrelated trade-offs: balancing the benefits and risks of obtaining the energy needed to cope with stressors, allocation of energy between life-history traits and stressor responses, and larger-scale escape from stressors in space or time via large-scale movement or dormancy. Finally, we outline how these trade-offs interactively affect fitness and qualitative ecological outcomes resulting from multiple stressors. Our framework suggests that explicitly considering animal behaviour should enrich our mechanistic understanding of stressor effects, help explain extensive context dependence observed in these effects, and highlight promising avenues for future empirical and theoretical research.

Understanding the mechanisms by which individual organisms respond and populations adapt to global climate change is a critical challenge. The role of plasticity and acclimation, within and across generations, may be essential given the pace of change. We investigated plasticity across generations and life stages in response to ocean acidification (OA), which poses a growing threat to both wild populations and the sustainable aquaculture of shellfish. Most studies of OA on shellfish focus on acute effects, and less is known regarding the longer term carryover effects that may manifest within or across generations. We assessed these longer term effects in red abalone ( Haliotis rufescens ) using a multi-generational split-brood experiment. We spawned adults raised in ambient conditions to create offspring that we then exposed to high pCO 2 (1180 μatm; simulating OA) or low pCO 2 (450 μatm; control or ambient conditions) during the first 3 months of life. We then allowed these animals to reach maturity in ambient common garden conditions for 4 years before returning the adults into high or low pCO 2 treatments for 11 months and measuring growth and reproductive potential. Early-life exposure to OA in the F1 generation decreased adult growth rate even after 5 years especially when abalone were re-exposed to OA as adults. Adult but not early-life exposure to OA negatively impacted fecundity. We then exposed the F2 offspring to high or low pCO 2 treatments for the first 3 months of life in a fully factorial, split-brood design. We found negative transgenerational effects of parental OA exposure on survival and growth of F2 offspring, in addition to significant direct effects of OA on F2 survival. These results show that the negative impacts of OA can last within and across generations, but that buffering against OA conditions at critical life-history windows can mitigate these effects.

Summary Conditional mutualisms involve costs and benefits that vary with environmental factors, but mechanisms driving these dynamics remain poorly understood. Scatterhoarder–plant interactions are a prime example of this phenomenon, as scatterhoarders can either increase or reduce plant recruitment depending on the balance between seed dispersal and predation. We explored factors that drive the magnitude of net benefits for plants in this interaction using a mathematical model, with parameter values based on European beech ( Fagus sylvatica ) and yellow‐necked mice ( Apodemus flavicollis ). We measured benefits as the percentage of germinating seeds, and examined how varying rodent survival (reflecting, e.g. changes in predation pressure), the rate of seed loss to other granivores, the abundance of alternative food resources, and changes in masting patterns affect the quality of mutualism. We found that increasing granivore abundance can degrade the quality of plant–scatterhoarder mutualism due to increased cache pilferage. Scatterhoarders are predicted to respond by increasing immediate consumption of gathered seeds, leading to higher costs and reduced benefits for plants. Thus, biotic changes that are detrimental to rodent populations can be beneficial for tree recruitment due to adaptive behavior of rodents. When scatterhoarder populations decline too drastically (< 5 individuals ha −1 ); however, tree recruitment may also suffer.