Human protection drives the emergence of a new coping style in animals.

Wild animals face novel environmental threats from human activities that may occur along a gradient of interactions with humans. Recent work has shown that merely living close to humans has major implications for a variety of antipredator traits and physiological responses. Here, we hypothesize that when human presence protects prey from their genuine predators (as sometimes seen in urban areas and at some tourist sites), this predator shield, followed by a process of habituation to humans, decouples commonly associated traits related to coping styles, which results in a new range of phenotypes. Such individuals are characterized by low aggressiveness and physiological stress responses, but have enhanced behavioral plasticity, boldness, and cognitive abilities. We refer to these individuals as "preactive," because their physiological and behavioral coping style falls outside the classical proactive/reactive coping styles. While there is some support for this new coping style, formal multivariate studies are required to investigate behavioral and physiological responses to anthropogenic activities.

Social control is associated with increased reproductive skew in a wild mammal.

In group-living species, reproductive variation among individuals of the same sex is widespread. By identifying the mechanisms underlying this reproductive skew, we gain fundamental insights into the evolution and maintenance of sociality. A common mechanism, social control, is typically studied by quantifying dominance, which is one of many attributes of sociality that describes how individuals exert influence on others and is an incomprehensive measure of social control as it accounts only for direct relationships. Here, we use the global reaching centrality (GRC), which quantifies the degree of hierarchy in a social network by accounting for both direct and indirect social relationships. Using a wild, free-living population of adult female yellow-bellied marmots (Marmota flaviventris), we found a positive relationship between the reproductive skew index and GRC: more despotic social groups have higher reproductive skew. The GRC was stronger predictor for skew than traditional measures of social control (i.e. dominance). This allows deeper insights into the diverse ways individuals control other group members' reproduction, a core component in the evolution of sociality. Future studies of skew across taxa may profit by using more comprehensive, network-based measures of social control.

Urban birds become less fearful following COVID-19 reopenings.

Following the COVID-19 pandemic, many people around the world stayed home, drastically altering human activity in cities. This exceptional moment provided researchers the opportunity to test how urban animals respond to human disturbance, in some cases testing fundamental questions on the mechanistic impact of urban behaviours on animal behaviour. However, at the end of this 'anthropause', human activity returned to cities. How might each of these strong shifts affect wildlife in the short and long term? We focused on fear response, a trait essential to tolerating urban life. We measured flight initiation distance-at both individual and population levels-for an urban bird before, during and after the anthropause to examine if birds experienced longer-term changes after a year and a half of lowered human presence. Dark-eyed juncos did not change fear levels during the anthropause, but they became drastically less fearful afterwards. These surprising and counterintuitive findings, made possible by following the behaviour of individuals over time, has led to a novel understanding that fear response can be driven by plasticity, yet not habituation-like processes. The pandemic-caused changes in human activity have shown that there is great complexity in how humans modify a behavioural trait fundamental to urban tolerance in animals.

Current and lagged climate affects phenology across diverse taxonomic groups.

The timing of life events (phenology) can be influenced by climate. Studies from around the world tell us that climate cues and species' responses can vary greatly. If variation in climate effects on phenology is strong within a single ecosystem, climate change could lead to ecological disruption, but detailed data from diverse taxa within a single ecosystem are rare. We collated first sighting and median activity within a high-elevation environment for plants, insects, birds, mammals and an amphibian across 45 years (1975-2020). We related 10 812 phenological events to climate data to determine the relative importance of climate effects on species' phenologies. We demonstrate significant variation in climate-phenology linkage across taxa in a single ecosystem. Both current and prior climate predicted changes in phenology. Taxa responded to some cues similarly, such as snowmelt date and spring temperatures; other cues affected phenology differently. For example, prior summer precipitation had no effect on most plants, delayed first activity of some insects, but advanced activity of the amphibian, some mammals, and birds. Comparing phenological responses of taxa at a single location, we find that important cues often differ among taxa, suggesting that changes to climate may disrupt synchrony of timing among taxa.

Individual life histories: neither slow nor fast, just diverse.

The slow-fast continuum is a commonly used framework to describe variation in life-history strategies across species. Individual life histories have also been assumed to follow a similar pattern, especially in the pace-of-life syndrome literature. However, whether a slow-fast continuum commonly explains life-history variation among individuals within a population remains unclear. Here, we formally tested for the presence of a slow-fast continuum of life histories both within populations and across species using detailed long-term individual-based demographic data for 17 bird and mammal species with markedly different life histories. We estimated adult lifespan, age at first reproduction, annual breeding frequency, and annual fecundity, and identified the main axes of life-history variation using principal component analyses. Across species, we retrieved the slow-fast continuum as the main axis of life-history variation. However, within populations, the patterns of individual life-history variation did not align with a slow-fast continuum in any species. Thus, a continuum ranking individuals from slow to fast living is unlikely to shape individual differences in life histories within populations. Rather, individual life-history variation is likely idiosyncratic across species, potentially because of processes such as stochasticity, density dependence, and individual differences in resource acquisition that affect species differently and generate non-generalizable patterns across species.

Foraging for foundations in decision neuroscience: insights from ethology.

Modern decision neuroscience offers a powerful and broad account of human behaviour using computational techniques that link psychological and neuroscientific approaches to the ways that individuals can generate near-optimal choices in complex controlled environments. However, until recently, relatively little attention has been paid to the extent to which the structure of experimental environments relates to natural scenarios, and the survival problems that individuals have evolved to solve. This situation not only risks leaving decision-theoretic accounts ungrounded but also makes various aspects of the solutions, such as hard-wired or Pavlovian policies, difficult to interpret in the natural world. Here, we suggest importing concepts, paradigms and approaches from the fields of ethology and behavioural ecology, which concentrate on the contextual and functional correlates of decisions made about foraging and escape and address these lacunae.

Evolutionary dynamics in the Anthropocene: Life history and intensity of human contact shape antipredator responses.

Humans profoundly impact landscapes, ecosystems, and animal behavior. In many cases, animals living near humans become tolerant of them and reduce antipredator responses. Yet, we still lack an understanding of the underlying evolutionary dynamics behind these shifts in traits that affect animal survival. Here, we used a phylogenetic meta-analysis to determine how the mean and variability in antipredator responses change as a function of the number of generations spent in contact with humans under 3 different contexts: urbanization, captivity, and domestication. We found that any contact with humans leads to a rapid reduction in mean antipredator responses as expected. Notably, the variance among individuals over time observed a short-term increase followed by a gradual decrease, significant for domesticated animals. This implies that intense human contact immediately releases animals from predation pressure and then imposes strong anthropogenic selection on traits. In addition, our results reveal that the loss of antipredator traits due to urbanization is similar to that of domestication but occurs 3 times more slowly. Furthermore, the rapid disappearance of antipredator traits was associated with 2 main life-history traits: foraging guild and whether the species was solitary or gregarious (i.e., group-living). For domesticated animals, this decrease in antipredator behavior was stronger for herbivores than for omnivores or carnivores and for solitary than for gregarious species. By contrast, the decrease in antipredator traits was stronger for gregarious, urbanized species, although this result is based mostly on birds. Our study offers 2 major insights on evolution in the Anthropocene: (1) changes in traits occur rapidly even under unintentional human "interventions" (i.e., urbanization) and (2) there are similarities between the selection pressures exerted by domestication and by urbanization. In all, such changes could affect animal survival in a predator-rich world, but through understanding evolutionary dynamics, we can better predict when and how exposure to humans modify these fitness-related traits.

Addressing prey naivety in native mammals by accelerating selection for antipredator traits.

Harnessing natural selection to improve conservation outcomes is a recent concept in ecology and evolutionary biology and a potentially powerful tool in species conservation. One possible application is the use of natural selection to improve antipredator responses of mammal species that are threatened by predation from novel predators. We investigated whether long-term exposure of an evolutionary naïve prey species to a novel predator would lead to phenotypic changes in a suite of physical and behavioral traits. We exposed a founder population of 353 burrowing bettongs (Bettongia lesueur) to feral cats (Felis catus) over 5 years and compared the physical and behavioral traits of this population (including offspring) to a control (non-predator exposed) population. We used selection analysis to investigate whether changes in the traits of bettongs were likely due to phenotypic plasticity or natural selection. We also quantified selection in both populations before and during major population crashes caused by drought (control) and high predation pressure (predator-exposed). Results showed that predator-exposed bettongs had longer flight initiation distances, larger hind feet, and larger heads than control bettongs. Trait divergence began soon after exposure and continued to intensify over time for flight initiation distance and hind foot length relative to control bettongs. Selection analysis found indicators of selection for larger hind feet and longer head length in predator-exposed populations. Results of a common garden experiment showed that the progeny of predator-exposed bettongs had larger feet than control bettongs. Results suggest that long-term, low-level exposure of naïve prey to novel predators can drive phenotypic changes that may assist with future conservation efforts.

Emergent social structure is typically not associated with survival in a facultatively social mammal.

For social animals, group social structure has important consequences for disease and information spread. While prior studies showed individual connectedness within a group has fitness consequences, less is known about the fitness consequences of group social structure for the individuals who comprise the group. Using a long-term dataset on a wild population of facultatively social yellow-bellied marmots (Marmota flaviventer), we showed social structure had largely no relationship with survival, suggesting consequences of individual social phenotypes may not scale to the group social phenotype. An observed relationship for winter survival suggests a potentially contrasting direction of selection between the group and previous research on the individual level; less social individuals, but individuals in more social groups experience greater winter survival. This work provides valuable insights into evolutionary implications across social phenotypic scales.

Older mothers produce more successful daughters.

Annual reproductive success and senescence patterns vary substantially among individuals in the wild. However, it is still seldom considered that senescence may not only affect an individual but also affect age-specific reproductive success in its offspring, generating transgenerational reproductive senescence. We used long-term data from wild yellow-bellied marmots (Marmota flaviventer) living in two different elevational environments to quantify age-specific reproductive success of daughters born to mothers differing in age. Contrary to prediction, daughters born to older mothers had greater annual reproductive success on average than daughters born to younger mothers, and this translated into greater lifetime reproductive success. However, in the favorable lower elevation environment, daughters born to older mothers also had greater age-specific decreases in annual reproductive success. In the harsher higher elevation environment on the other hand, daughters born to older mothers tended to die before reaching ages at which such senescent decreases could be observed. Our study highlights the importance of incorporating environment-specific transgenerational parent age effects on adult offspring age-specific life-history traits to fully understand the substantial variation observed in senescence patterns in wild populations.

Contrasting effects of climate change on seasonal survival of a hibernating mammal.

Seasonal environmental conditions shape the behavior and life history of virtually all organisms. Climate change is modifying these seasonal environmental conditions, which threatens to disrupt population dynamics. It is conceivable that climatic changes may be beneficial in one season but result in detrimental conditions in another because life-history strategies vary between these time periods. We analyzed the temporal trends in seasonal survival of yellow-bellied marmots (Marmota flaviventer) and explored the environmental drivers using a 40-y dataset from the Colorado Rocky Mountains (USA). Trends in survival revealed divergent seasonal patterns, which were similar across age-classes. Marmot survival declined during winter but generally increased during summer. Interestingly, different environmental factors appeared to drive survival trends across age-classes. Winter survival was largely driven by conditions during the preceding summer and the effect of continued climate change was likely to be mainly negative, whereas the likely outcome of continued climate change on summer survival was generally positive. This study illustrates that seasonal demographic responses need disentangling to accurately forecast the impacts of climate change on animal population dynamics.

Welcome to the Pyrocene: Animal survival in the age of megafire.

Planet Earth is entering the age of megafire, pushing ecosystems to their limits and beyond. While fire causes mortality of animals across vast portions of the globe, scientists are only beginning to consider fire as an evolutionary force in animal ecology. Here, we generate a series of hypotheses regarding animal responses to fire by adopting insights from the predator-prey literature. Fire is a lethal threat; thus, there is likely strong selection for animals to recognize the olfactory, auditory, and visual cues of fire, and deploy fire avoidance behaviours that maximize survival probability. If fire defences are costly, it follows that intraspecific variation in fire avoidance behaviours should correspond with variation in fire behaviour and regimes. Species and populations inhabiting ecosystems that rarely experience fire may lack these traits, placing 'fire naive' populations and species at enhanced extinction risk as the distribution of fire extends into new ecosystem types. We outline a research agenda to understand behavioural responses to fire and to identify conservation interventions that could be used to overcome fire naivety.

Social position indirectly influences the traits yellow-bellied marmots use to solve problems.

Animals adapt to changing environments by behaving flexibly when solving problems. Traits, such as sex and age, and specifically behavioral traits like persistence-the amount of time spent attempting to solve a problem, are positively associated with successful problem-solving. However, individuals face social pressures, such as aggression, which may directly alter an individual's behavior or interact with sex or age, when they attempt to problem-solve. We examined the direct and indirect effects of social position and individual behavioral traits on solving a novel puzzle box in facultatively social yellow-bellied marmots (Marmota flaviventer), using both generalized linear mixed models and confirmatory path analysis. We found strong support that marmots who used a diversity of behaviors were more successful problem-solvers and weak support that those who received more aggression were less successful. Additionally, marmots who received more aggression were less behaviorally diverse, less behaviorally selective and less persistent while trying to open the puzzle box. Thus, we show that aggression indirectly decreases problem-solving success by acting on the behavioral traits that an individual uses. We conclude that specific social relationships, including the type of interaction and whether they are recipients or initiators, influences the ways in which an individual interacts with cognitive tests and should be considered in analysis of individual problem-solving.

Selective consumption of macroalgal species by herbivorous fishes suggests reduced functional complementarity on a fringing reef in Moorea, French Polynesia.

Worldwide, many coral reefs are at risk of shifting to degraded algal-dominated states, due to compromised ecological conditions. Functional diversity of herbivorous fishes maintains coral reef health and promotes reef resilience to disturbances. Given previous evidence, it appears the functional roles of herbivorous fishes differ across geographical locations, indicating a need for further assessment of macroalgal consumption by herbivorous fishes. We assessed functional diversity by examining foraging behavior of herbivorous fish species on macroalgae on a fringing reef in Moorea, French Polynesia. We video-recorded choice experiments containing seven common macroalgae and used Strauss' linear resource selection index to determine macroalgal selectivities. We used cluster analysis to identify any distinct groups within herbivorous fish species, given the macroalgal species they targeted, and fitted generalized linear mixed-effects models to identify factors that best predicted the number of bites taken on macroalgae. Seven species from 3 fish families/tribes took a total of 956 bites. Fish species differed in their selectivity with some species (Naso lituratus, N. unicornis, Calatomus carolinus) strongly preferring one or two macroalgal species, while other fish species (Acanthurus nigrofuscus, Ctenochaetus striatus, Chlorurus sordidus, Balistapus undulatus) were less selective. This resulted in fish species forming two clusters. Only 3 of 7 macroalgae were preferred by any fish species, with two fish species both preferring the same two macroalgae. The limited differences in fish species' preferences for different macroalgae suggests limited functional complementarity. Two models (macroalgal species identity+fish functional group, macroalgal species identity+fish species) best predicted the number of bites taken on macroalgae compared to models incorporating only a single explanatory factor or fish family. In the context of this Moorean fringing reef, there is greater functional redundancy than complementarity of herbivorous fishes consuming macroalgae, and the fishes grouped together according to their relative selectivity. We observed fish species that are not classified as browsers consuming macroalgae, suggesting diets of herbivorous fishes may be broader than previously thought. Finally, we observed macroalgal selectivities and consumption that differed from previous studies for the same fish species. Our results contribute to the understanding of functional diversity of herbivorous fishes across coral reefs, and also highlight the need for additional research to further elucidate the role of context and functional diversity of herbivorous fishes consuming macroalgae on coral reefs.

Assessing seasonal demographic covariation to understand environmental-change impacts on a hibernating mammal.

Natural populations are exposed to seasonal variation in environmental factors that simultaneously affect several demographic rates (survival, development and reproduction). The resulting covariation in these rates determines population dynamics, but accounting for its numerous biotic and abiotic drivers is a significant challenge. Here, we use a factor-analytic approach to capture partially unobserved drivers of seasonal population dynamics. We use 40 years of individual-based demography from yellow-bellied marmots (Marmota flaviventer) to fit and project population models that account for seasonal demographic covariation using a latent variable. We show that this latent variable, by producing positive covariation among winter demographic rates, depicts a measure of environmental quality. Simultaneously, negative responses of winter survival and reproductive-status change to declining environmental quality result in a higher risk of population quasi-extinction, regardless of summer demography where recruitment takes place. We demonstrate how complex environmental processes can be summarized to understand population persistence in seasonal environments.

Safety Cues Can Give Prey More Valuable Information Than Danger Cues.

The ability of prey to assess predation risk is fundamental to their success. It is routinely assumed that predator cues do not vary in reliability across levels of predation risk. We propose that cues can differ in how precisely they indicate different levels of predation risk. What we call danger cues precisely indicate high risk levels, while safety cues precisely indicate low risk levels. Using optimality modeling, we find that prey fitness is increased when prey pay more attention to safety cues than to danger cues. This fitness advantage is greater when prey need to protect assets, predators are more dangerous, or predation risk increases at an accelerating rate with prey foraging efforts. Each of these conditions lead to prey foraging less when estimated predation risk is higher. Danger cues have less value than safety cues because they give precise information about risk when it is high, but prey behavior varies little when risk is high. Safety cues give precise information about levels of risk where prey behavior varies. These results highlight how our fascination with predators may have biased the way that we study predator-prey interactions and focused too exclusively on cues that clearly indicate the presence of predator rather than cues that clearly indicate their absence.

Correlates of maternal glucocorticoid levels in a socially flexible rodent.

While it is generally accepted that social isolation has detrimental effects on social species, little is known about the importance of social interactions in less social species-particularly for wild reproductive females. We studied socially-flexible yellow-bellied marmots (Marmota flaviventer) and asked whether features of the social environment are associated with maternal fecal glucocorticoid metabolite (FGM) concentrations. Since changes in maternal baseline glucocorticoids may have positive or negative consequences for offspring fitness, we were also interested in estimating their relationship with measures of reproductive success. We fitted generalized linear mixed effects models to a dataset including maternal FGM measurements, social network metrics, maternal/alloparental care, and pup FGM and survival. Agonistic interactions were positively associated with maternal FGM levels, while mothers that engaged in relatively more affiliative interactions had reduced FGM levels when living in environments with low predator pressure. Pups associated with mothers exhibiting high FGM levels had low annual survival rates, received less maternal/alloparental care and had higher FGM levels. Interestingly, offspring from mothers with high FGM levels were more likely to survive the summer when born in small litters. In sum, social interactions likely influence and are influenced by glucocorticoid levels of facultatively social females. Potential benefits of social bonds may be context-specific, and agonistic interactions may be tightly correlated with fitness. Female marmots exhibiting high FGM levels had overall low reproductive success, which is predicted by the cort-fitness hypothesis. However, under adverse conditions, offspring summer survival can be maximized if pups are born in small litters.

Downsizing for downtown: Limb lengths, toe lengths, and scale counts decrease with urbanization in western fence lizards (Sceloporus occidentalis).

Urbanization induced habitat loss and alteration causes significant challenges for the survival of many species. Identifying how species respond to urbanization can yield insights for the conservation of wildlife, but research on reptiles has been narrowly-focused. We compared morphology among four populations of western fence lizards (Sceloporus occidentalis) to determine whether a common native species affected by urbanization exhibits morphological differences consistent with habitat use. We quantified habitat differences across four sites in Los Angeles County, California, USA that varied in level of urbanization, measured how lizards used microhabitats, and assessed variation in body size, limb length, toe length, and scalation of lizards collected from each site. Urban and suburban populations of fence lizards mostly used human-made substrates while lizards from more natural areas mostly used natural woody substrates. Lizards from the most urban site also exhibited the widest breadth of substrates used, indicating that urban sites might offer more variable microhabitats. Urban lizards had reduced limb lengths and toe lengths consistent with how they used microhabitats and other habitat characteristics (e.g., percent impervious surface cover). Urban lizards also had fewer dorsal scales, which might be associated with changes in ambient temperature (e.g., urban heat island effect), given that lizards with fewer and larger scales typically have reduced evaporative water loss. Our results uniquely differ from past studies on lizard responses to urbanization, indicating that work on diverse taxa is necessary to assess the potential varied pathways of morphological adaptations to urban environments.

Transient LTRE analysis reveals the demographic and trait-mediated processes that buffer population growth.

Temporal variation in environmental conditions affects population growth directly via its impact on vital rates, and indirectly through induced variation in demographic structure and phenotypic trait distributions. We currently know very little about how these processes jointly mediate population responses to their environment. To address this gap, we develop a general transient life table response experiment (LTRE) which partitions the contributions to population growth arising from variation in (1) survival and reproduction, (2) demographic structure, (3) trait values and (4) climatic drivers. We apply the LTRE to a population of yellow-bellied marmots (Marmota flaviventer) to demonstrate the impact of demographic and trait-mediated processes. Our analysis provides a new perspective on demographic buffering, which may be a more subtle phenomena than is currently assumed. The new LTRE framework presents opportunities to improve our understanding of how trait variation influences population dynamics and adaptation in stochastic environments.

Strong social relationships are associated with decreased longevity in a facultatively social mammal.

Humans in strong social relationships are more likely to live longer because social relationships may buffer stressors and thus have protective effects. However, a shortcoming of human studies is that they often rely on self-reporting of these relationships. By contrast, observational studies of non-human animals permit detailed analyses of the specific nature of social relationships. Thus, discoveries that some social animals live longer and healthier lives if they are involved in social grooming, forage together or have more affiliative associates emphasizes the potential importance of social relationships on health and longevity. Previous studies have focused on the impact of social metrics on longevity in obligately social species. However, if sociality indeed has a key role in longevity, we might expect that affiliative relationships should also influence longevity in less social species. We focused on socially flexible yellow-bellied marmots (Marmota flaviventer) and asked whether female longevity covaries with the specific nature of social relationships. We quantified social relationships with social network statistics that were based on affiliative interactions, and then estimated the correlation between longevity and sociality using bivariate models. We found a significant negative phenotypic correlation between affiliative social relationship strength and longevity; marmots with greater degree, closeness and those with a greater negative average shortest path length died at younger ages. We conclude that sociality plays an important role in longevity, but how it does so may depend on whether a species is obligately or facultatively social.