Environmental harshness does not affect the propensity for social learning in great tits, Parus major.

According to the harsh environment hypothesis, natural selection should favour cognitive mechanisms to overcome environmental challenges. Tests of this hypothesis to date have largely focused on asocial learning and memory, thus failing to account for the spread of information via social means. Tests in specialized food-hoarding birds have shown strong support for the effects of environmental harshness on both asocial and social learning. Whether the hypothesis applies to non-specialist foraging species remains largely unexplored. We evaluated the relative importance of social learning across a known harshness gradient by testing generalist great tits, Parus major, from high (harsh)- and low (mild)-elevation populations in two social learning tasks. We showed that individuals use social learning to find food in both colour-associative and spatial foraging tasks and that individuals differed consistently in their use of social learning. However, we did not detect a difference in the use or speed of implementing socially observed information across the elevational gradient. Our results do not support predictions of the harsh environment hypothesis suggesting that context-dependent costs and benefits as well as plasticity in the use of social information may play an important role in the use of social learning across environments. Finally, this study adds to the accumulating evidence that the harsh environment hypothesis appears to have more pronounced effects on specialists compared to generalist species.

Texas field crickets (Gryllus texensis) use visual cues to place learn but perform poorly when intra- and extra-maze cues conflict.

Central place foraging field crickets are an ideal system for studying the adaptive value of learning and memory, but more research is needed on ecologically relevant cognition in these invertebrates. Here, we test the visuospatial place learning of Texas field crickets (Gryllus texensis) in a radial arm maze. Our study expands previous work on G. texensis cognition for accuracy measures and extends our previous findings on females to both sexes. Additionally, our study examines whether crickets use intra- or extra-maze cues to locate a food reward using a maze rotation that puts the cues in conflict. We found that male and female crickets improved performance over trials when measured by accuracy variables but not latency variables. Thigmotaxis negatively impacted performance in both sexes. In a reward-absent trial, both male and female crickets demonstrated place memory. When intra- and extra-maze cues conflicted during a rotation trial, crickets' performance was not better than chance. Our rotation results suggest that crickets may experience reciprocal overshadowing of conflicting cues - a result most often seen in other taxa with conflicting multi-modal cues. We conclude that crickets do not rely solely on: (1) a single-cue association, (2) route-following, or (3) their own scent cues to navigate the maze. Instead, male and female Texas field crickets seem to learn the location of the reward using a combination of proximal and distal cues. The possibility to test large numbers of wild-caught or laboratory-reared individuals opens the door to future investigations on the evolutionary ecology of visuospatial learning in these invertebrates.

Cognitive flexibility in the wild: Individual differences in reversal learning are explained primarily by proactive interference, not by sampling strategies, in two passerine bird species.

Behavioural flexibility allows animals to adjust to changes in their environment. Although the cognitive processes that explain flexibility have been relatively well studied in psychology, this is less true for animals in the wild. Here we use data collected automatically during self-administered discrimination-learning trials for two passerine species, and during four phases (habituation, initial learning, first reversal and second reversal) in order to decompose sources of consistent among-individual differences in reversal learning, a commonly used measure for cognitive flexibility. First, we found that, as expected, proactive interference was significantly repeatable and had a negative effect on reversal learning, confirming that individuals with poor ability to inhibit returning to a previously rewarded feeder were also slower to reversal learn. Second, to our knowledge for the first time in a natural population, we examined how sampling of non-rewarding options post-learning affected reversal-learning performance. Sampling quantity was moderately repeatable in blue tits but not great tits; sampling bias, the variance in the proportion of visits to each non-rewarded feeder, was not repeatable for either species. Sampling behaviour did not predict variation in reversal-learning speed to any significant extent. Finally, the repeatability of reversal learning was explained almost entirely by proactive interference for blue tits; in great tits, the effects of proactive interference and sampling bias on the repeatability of reversal learning were indistinguishable. Our results highlight the value of proactive interference as a more direct measurement of cognitive flexibility and shed light on how animals respond to changes in their environment.

Cognition and covariance in the producer-scrounger game.

The producer-scrounger game is a key element of foraging ecology in many systems. Producing and scrounging typically covary negatively, but partitioning this covariance into contributions of individual plasticity and consistent between individual differences is key to understanding population-level consequences of foraging strategies. Furthermore, little is known about the role cognition plays in the producer-scrounger game. We investigated the role of cognition in these alternative foraging tactics in wild mixed-species flocks of great tits and blue tits, using a production learning task in which we measured individuals' speed of learning to visit the single feeder in an array that would provide them with a food reward. We also quantified the proportion of individuals' feeds that were scrounges ('proportion scrounged'); scrounging was possible if individuals visited immediately after a previous rewarded visitor. Three learning experiments-initial and two reversal learning-enabled us to estimate the repeatability and covariance of each foraging behaviour. First, we examined whether individuals learned to improve their scrounging success (i.e. whether they obtained food by scrounging when there was an opportunity to do so). Second, we quantified the repeatability of proportion scrounged, and asked whether proportion scrounged affected production learning speed among individuals. Third, we used multivariate analyses to partition within- and among-individual components of covariance between proportion scrounged and production learning speed. Individuals improved their scrounging success over time. Birds with a greater proportion scrounged took longer to learn their own rewarding feeder. Although multivariate analyses showed that covariance between proportion scrounged and learning speed was driven primarily by within-individual variation, that is, by behavioural plasticity, among-individual differences also played a role for blue tits. This is the first demonstration of a cognitive trait influencing producing and scrounging in the same wild system, highlighting the importance of cognition in the use of alternative resource acquisition tactics. The results of our covariance analyses suggest the potential for genetic differences in allocation to alternative foraging tactics, which are likely species- and system-dependent. They also point to the need to control for different foraging tactics when studying individual cognition in the wild.

Great tits who remember more accurately have difficulty forgetting, but variation is not driven by environmental harshness.

The causes of individual variation in memory are poorly understood in wild animals. Harsh environments with sparse or rapidly changing food resources are hypothesized to favour more accurate spatial memory to allow animals to return to previously visited patches when current patches are depleted. A potential cost of more accurate spatial memory is proactive interference, where accurate memories block the formation of new memories. This relationship between spatial memory, proactive interference, and harsh environments has only been studied in scatter-hoarding animals. We compare spatial memory accuracy and proactive interference performance of non-scatter hoarding great tits (Parus major) from high and low elevations where harshness increases with elevation. In contrast to studies of scatter-hoarders, we did not find a significant difference between high and low elevation birds in their spatial memory accuracy or proactive interference performance. Using a variance partitioning approach, we report the first among-individual trade-off between spatial memory and proactive interference, uncovering variation in memory at the individual level where selection may act. Although we have no evidence of harsh habitats affecting spatial memory, our results suggest that if elevation produced differences in spatial memory between elevations, we could see concurrent changes in how quickly birds can forget.

How general is cognitive ability in non-human animals? A meta-analytical and multi-level reanalysis approach.

General intelligence has been a topic of high interest for over a century. Traditionally, research on general intelligence was based on principal component analyses and other dimensionality reduction approaches. The advent of high-speed computing has provided alternative statistical tools that have been used to test predictions of human general intelligence. In comparison, research on general intelligence in non-human animals is in its infancy and still relies mostly on factor-analytical procedures. Here, we argue that dimensionality reduction, when incorrectly applied, can lead to spurious results and limit our understanding of ecological and evolutionary causes of variation in animal cognition. Using a meta-analytical approach, we show, based on 555 bivariate correlations, that the average correlation among cognitive abilities is low (r = 0.185; 95% CI: 0.087-0.287), suggesting relatively weak support for general intelligence in animals. We then use a case study with relatedness (genetic) data to demonstrate how analysing traits using mixed models, without dimensionality reduction, provides new insights into the structure of phenotypic variance among cognitive traits, and uncovers genetic associations that would be hidden otherwise. We hope this article will stimulate the use of alternative tools in the study of cognition and its evolution in animals.

The impact of learning opportunities on the development of learning and decision-making: an experiment with passerine birds.

Developmental context has been shown to influence learning abilities later in life, namely through experiments with nutritional and/or environmental constraints (i.e. lack of enrichment). However, little is known about the extent to which opportunities for learning affect the development of animal cognition, even though such opportunities are known to influence human cognitive development. We exposed young zebra finches (Taenopygia guttata) (n = 26) to one of three experimental conditions, i.e. an environment where (i) colour cues reliably predicted the presence of food (associative learning), (ii) a combination of two-colour cues reliably predicted the presence of food (conditional learning), or (iii) colour cues were non-informative (control). After conducting two different discrimination tasks, our results showed that experience with predictive cues can cause increased choice accuracy and decision-making speed. Our first learning task showed that individuals in the associative learning treatment outperformed the control treatment, while task 2 showed that individuals in the conditional learning treatment had shorter latencies when making choices compared with the control treatment. We found no support for a speed-accuracy trade-off. This dataset provides a rare longitudinal and experimental examination of the effect of predictive versus non-predictive cues during development on the cognition of adult animals. This article is part of the theme issue 'Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals'.

Characterizing innovators: Ecological and individual predictors of problem-solving performance.

Behavioural innovation, the use of new behaviours or existing ones in novel contexts, can have important ecological and evolutionary consequences for animals. An understanding of these consequences would be incomplete without considering the traits that predispose certain individuals to exhibit innovative behaviour. Several individual and ecological variables are hypothesized to affect innovativeness, but empirical studies show mixed results. We examined the effects of dominance rank, exploratory personality, and urbanisation on the innovativeness of wild-caught black-capped chickadees using a survival analysis of their performance in two problem-solving tasks. Additionally, we provide one of the first investigations of the predictors of persistence in a problem-solving context. For lever pulling, we found a trend for dominants to outperform subordinates, particularly in rural birds, which did not align with predictions from the necessity drives innovation hypothesis. When examining possible explanations for this trend we found that older chickadees outperformed younger birds. This follow-up analysis also revealed a positive effect of exploratory personality on the lever-pulling performance of chickadees. Our results suggest that experience may foster innovation in certain circumstances, for instance via the application of previously-acquired information or skills to a novel problem. As we found different predictors for both tasks, this suggests that task characteristics influence the innovative propensity of individuals, and that their effects should be investigated experimentally.

Food caching in city birds: urbanization and exploration do not predict spatial memory in scatter hoarders.

Urbanization has been shown to affect the physiological, morphological, and behavioral traits of animals, but it is less clear how cognitive traits are affected. Urban habitats contain artificial food sources, such as bird feeders that are known to impact foraging behaviors. As of yet, however, it is not well known whether urbanization and the abundance of supplemental food during the winter affect caching behaviors and spatial memory in scatter hoarders. We aim to compare caching intensity and spatial memory performance along an urban gradient to determine (i) whether individuals from more urbanized sites cache less frequently and perform less accurately on a spatial memory task, and (ii) for the first time in individual scatter hoarders, whether slower explorers perform more accurately than faster explorers on a spatial memory task. We assessed food caching, exploration of a novel environment, and spatial memory performance of wild-caught black-capped chickadees (Poecile atricapillus; N = 95) from 14 sites along an urban gradient. Although the individuals that cached most in captivity were all from less urbanized sites, we found no clear evidence that caching intensity and spatial memory accuracy differed along an urban gradient. At the individual level, we found no significant relationship between spatial memory performance and exploration score. However, individuals that performed more accurately on the spatial task also tended to cache more, pointing to a specialization of spatial memory in scatter hoarders that could occur at the level of the individual, in addition to the previously documented specialization at the population and species levels.

Urbanization and the temporal patterns of social networks and group foraging behaviors.

Urbanization causes dramatic and rapid changes to natural environments, which can lead the animals inhabiting these habitats to adjust their behavioral responses. For social animals, urbanized environments may alter group social dynamics through modification of the external environment (e.g., resource distribution). This might lead to changes in how individuals associate or engage in group behaviors, which could alter the stability and characteristics of social groups. However, the potential impacts of urban habitat use, and of habitat characteristics in general, on the nature and stability of social associations remain poorly understood. Here, we quantify social networks and dynamics of group foraging behaviors of black-capped chickadees (N = 82, Poecile atricapillus), at four urban and four rural sites weekly throughout the nonbreeding season using feeders with radio frequency identification of individual birds. Because anthropogenic food sources in urban habitats (e.g., bird feeders) provide abundant and reliable resources, we predicted that social foraging associations may be of less value in urban groups, and thus would be less consistent than in rural groups. Additionally, decreased variability of food resources in urban habitats could lead to more predictable foraging patterns (group size, foraging duration, and the distribution of foraging events) in contrast to rural habitats. Networks were found to be highly consistent through time in both urban and rural habitats. No significant difference was found in the temporal clumping of foraging events between habitats. However, as predicted, the repeatability of the clumping of foraging events in time was significantly higher in urban than rural habitats. Our results suggest that individuals living in urban areas have more consistent foraging behaviors throughout the nonbreeding season, whereas rural individuals adjust their tactics due to less predictable foraging conditions. This first examination of habitat-related differences in the characteristics and consistency of social networks along an urbanization gradient suggests that anthropic habitat use results in subtle modifications in social foraging patterns. Future studies should examine potential implications of these differences for variation in predation risk, energy intake, and information flow.

Energy metabolism and personality in wild-caught fall field crickets.

Standard metabolic rate (SMR) is known to be highly variable across levels of biological organisation (e.g., species, populations, among individuals, within individuals). Some of the variation in SMR can be attributed to factors such as diet, temperature, and body mass, yet much of the residual variation in SMR remains unexplained. Intuitively, we can expect SMR to co-vary with "personality", but the rapidly accumulating empirical evidence on this topic remains equivocal. The goal of this study was to test for a link between SMR and a behavioural syndrome at the among-individual level in wild-caught fall field crickets (Gryllus pennsylvanicus). Paired measurements of SMR and two behavioural traits were repeatedly taken over a two-month period, thus allowing to estimate the among-individual correlations (rind) separately from the residual (within-individual) correlations. The two behavioural traits (latency to exit a refuge in a novel environment and "freezing" time following a stressful stimulus) were significantly and moderately repeatable and were found to be part of a syndrome, as indicated by a strong and positive among-individual correlation (rind = 0.82 ±â€¯0.27). Yet, only latency to exit was significantly and positively correlated with SMR (rind = 0.45 ±â€¯0.21), suggesting that the link between boldness and SMR may be driven by individual differences in responses to novelty and not to simulated predatory cues. Since we found that bold individuals (short latency to exit) consistently had a lower SMR than shy individuals (long latency to exit), our results go against the pace-of-life syndrome hypothesis. Instead, our results suggest the presence of constrains in the energy budgets of crickets, which generated an allocation trade-off between energy spent on physical activity vs. maintenance costs (SMR).

Measuring and understanding individual differences in cognition.

Individuals vary in their cognitive performance. While this variation forms the foundation of the study of human psychometrics, its broader importance is only recently being recognized. Explicitly acknowledging this individual variation found in both humans and non-human animals provides a novel opportunity to understand the mechanisms, development and evolution of cognition. The papers in this special issue highlight the growing emphasis on individual cognitive differences from fields as diverse as neurobiology, experimental psychology and evolutionary biology. Here, we synthesize this body of work. We consider the distinct challenges in quantifying individual differences in cognition and provide concrete methodological recommendations. In particular, future studies would benefit from using multiple task variants to ensure they target specific, clearly defined cognitive traits and from conducting repeated testing to assess individual consistency. We then consider how neural, genetic, developmental and behavioural factors may generate individual differences in cognition. Finally, we discuss the potential fitness consequences of individual cognitive variation and place these into an evolutionary framework with testable hypotheses. We intend for this special issue to stimulate researchers to position individual variation at the centre of the cognitive sciences.This article is part of the theme issue 'Causes and consequences of individual differences in cognitive abilities'.

Reduced reproductive performance associated with warmer ambient temperatures during incubation in a winter-breeding, food-storing passerine.

Timing of reproduction can influence individual fitness whereby early breeders tend to have higher reproductive success than late breeders. However, the fitness consequences of timing of breeding may also be influenced by environmental conditions after the commencement of breeding. We tested whether ambient temperatures during the incubation and early nestling periods modulated the effect of laying date on brood size and dominant juvenile survival in gray jays (Perisoreus canadensis), a sedentary boreal species whose late winter nesting depends, in part, on caches of perishable food. Previous evidence has suggested that warmer temperatures degrade the quality of these food hoards, and we asked whether warmer ambient temperatures during the incubation and early nestling periods would be associated with smaller brood sizes and lower summer survival of dominant juveniles. We used 38 years of data from a range-edge population of gray jays in Algonquin Provincial Park, Ontario, where the population has declined over 50% since the study began. Consistent with the "hoard-rot" hypothesis, we found that cold temperatures during incubation were associated with larger brood sizes in later breeding attempts, but temperatures had little effect on brood size for females breeding early in the season. This is the first evidence that laying date and temperature during incubation interactively influence brood size in any bird species. We did not find evidence that ambient temperatures during the incubation period or early part of the nestling period influenced summer survival of dominant juveniles. Our findings provide evidence that warming temperatures are associated with some aspects of reduced reproductive performance in a species that is reliant on cold temperatures to store perishable food caches, some of which are later consumed during the reproductive period.

Animal and human innovation: novel problems and novel solutions.

This theme issue explores how and why behavioural innovation occurs, and the consequences of innovation for individuals, groups and populations. A vast literature on human innovation exists, from the development of problem-solving in children, to the evolution of technology, to the cultural systems supporting innovation. A more recent development is a growing literature on animal innovation, which has demonstrated links between innovation and personality traits, cognitive traits, neural measures, changing conditions, and the current state of the social and physical environment. Here, we introduce these fields, define key terms and discuss the potential for fruitful exchange between the diverse fields researching innovation. Comparisons of innovation between human and non-human animals provide opportunities, but also pitfalls. We also summarize some key findings specifying the circumstances in which innovation occurs, discussing factors such as the intrinsic nature of innovative individuals and the environmental and socio-ecological conditions that promote innovation, such as necessity, opportunity and free resources. We also highlight key controversies, including the relationship between innovation and intelligence, and the notion of innovativeness as an individual-level trait. Finally, we discuss current research methods and suggest some novel approaches that could fruitfully be deployed.

Environmental and genetic determinants of innovativeness in a natural population of birds.

Much of the evidence for the idea that individuals differ in their propensity to innovate and solve new problems has come from studies on captive primates. Increasingly, behavioural ecologists are studying innovativeness in wild populations, and uncovering links with functional behaviour and fitness-related traits. The relative importance of genetic and environmental factors in driving this variation, however, remains unknown. Here, we present the results of the first large-scale study to examine a range of causal factors underlying innovative problem-solving performance (PSP) among 831 great tits (Parus major) temporarily taken into captivity. Analyses show that PSP in this population: (i) was linked to a variety of individual factors, including age, personality and natal origin (immigrant or local-born); (ii) was influenced by natal environment, because individuals had a lower PSP when born in poor-quality habitat, or where local population density was high, leading to cohort effects. Links with many of the individual and environmental factors were present only in some years. In addition, PSP (iii) had little or no measurable heritability, as estimated by a Bayesian animal model; and (iv) was not influenced by maternal effects. Despite previous reports of links between PSP and a range of functional traits in this population, the analyses here suggest that innovativeness had weak if any evolutionary potential. Instead most individual variation was caused by phenotypic plasticity driven by links with other behavioural traits and by environmentally mediated developmental stress. Heritability estimates are population, time and context specific, however, and more studies are needed to determine the generality of these effects. Our results shed light on the causes of innovativeness within populations, and add to the debate on the relative importance of genetic and environmental factors in driving phenotypic variation within populations.

Studying the evolutionary ecology of cognition in the wild: a review of practical and conceptual challenges.

Cognition is defined as the processes by which animals collect, retain and use information from their environment to guide their behaviour. Thus cognition is essential in a wide range of behaviours, including foraging, avoiding predators and mating. Despite this pivotal role, the evolutionary processes shaping variation in cognitive performance among individuals in wild populations remain very poorly understood. Selection experiments in captivity suggest that cognitive traits can have substantial heritability and can undergo rapid evolution. However only a handful of studies have attempted to explore how cognition influences life-history variation and fitness in the wild, and direct evidence for the action of natural or sexual selection on cognition is still lacking, reasons for which are diverse. Here we review the current literature with a view to: (i) highlighting the key practical and conceptual challenges faced by the field; (ii) describing how to define and measure cognitive traits in natural populations, and suggesting which species, populations and cognitive traits might be examined to greatest effect; emphasis is placed on selecting traits that are linked to functional behaviour; (iii) discussing how to deal with confounding factors such as personality and motivation in field as well as captive studies; (iv) describing how to measure and interpret relationships between cognitive performance, functional behaviour and fitness, offering some suggestions as to when and what kind of selection might be predicted; and (v) showing how an evolutionary ecological framework, more generally, along with innovative technologies has the potential to revolutionise the study of cognition in the wild. We conclude that the evolutionary ecology of cognition in wild populations is a rapidly expanding interdisciplinary field providing many opportunities for advancing the understanding of how cognitive abilities have evolved.

Taking the Operant Paradigm into the Field: Associative Learning in Wild Great Tits.

Associative learning is essential for resource acquisition, predator avoidance and reproduction in a wide diversity of species, and is therefore a key target for evolutionary and comparative cognition research. Automated operant devices can greatly enhance the study of associative learning and yet their use has been mainly restricted to laboratory conditions. We developed a portable, weatherproof, battery-operated operant device and conducted the first fully automated colour-associative learning experiment using free-ranging individuals in the wild. We used the device to run a colour discrimination task in a monitored population of tits (Paridae). Over two winter months, 80 individuals from four species recorded a total of 5,128 trials. Great tits (Parus major) were more likely than other species to visit the devices and engage in trials, but there were no sex or personality biases in the sample of great tits landing at the devices and registering key pecks. Juveniles were more likely than adults to visit the devices and to register trials. Individuals that were successful at solving a novel technical problem in captivity (lever-pulling) learned faster than non-solvers when at the operant devices in the wild, suggesting cross-contextual consistency in learning performance in very different tasks. There was no significant effect of personality or sex on learning rate, but juveniles' choice accuracy tended to improve at a faster rate than adults. We discuss how customisable automated operant devices, such as the one described here, could prove to be a powerful tool in evolutionary ecology studies of cognitive traits, especially among inquisitive species such as great tits.

The evolution of cognition in natural populations.

Individual differences in cognitive abilities have been described in a range of species, but do they impact survival or reproduction? Several recent studies report links between putative cognitive and reproductive traits in avian systems. Whether or when selection should occur in the wild is becoming an exciting avenue of research.

Experimentally induced innovations lead to persistent culture via conformity in wild birds.

In human societies, cultural norms arise when behaviours are transmitted through social networks via high-fidelity social learning. However, a paucity of experimental studies has meant that there is no comparable understanding of the process by which socially transmitted behaviours might spread and persist in animal populations. Here we show experimental evidence of the establishment of foraging traditions in a wild bird population. We introduced alternative novel foraging techniques into replicated wild sub-populations of great tits (Parus major) and used automated tracking to map the diffusion, establishment and long-term persistence of the seeded innovations. Furthermore, we used social network analysis to examine the social factors that influenced diffusion dynamics. From only two trained birds in each sub-population, the information spread rapidly through social network ties, to reach an average of 75% of individuals, with a total of 414 knowledgeable individuals performing 57,909 solutions over all replicates. The sub-populations were heavily biased towards using the technique that was originally introduced, resulting in established local traditions that were stable over two generations, despite a high population turnover. Finally, we demonstrate a strong effect of social conformity, with individuals disproportionately adopting the most frequent local variant when first acquiring an innovation, and continuing to favour social information over personal information. Cultural conformity is thought to be a key factor in the evolution of complex culture in humans. In providing the first experimental demonstration of conformity in a wild non-primate, and of cultural norms in foraging techniques in any wild animal, our results suggest a much broader taxonomic occurrence of such an apparently complex cultural behaviour.

Cognitive ability influences reproductive life history variation in the wild.

Cognition has been studied intensively for several decades, but the evolutionary processes that shape individual variation in cognitive traits remain elusive [1-3]. For instance, the strength of selection on a cognitive trait has never been estimated in a natural population, and the possibility that positive links with life history variation [1-5] are mitigated by costs [6] or confounded by ecological factors remains unexplored in the wild. We assessed novel problem-solving performance in 468 wild great tits Parus major temporarily taken into captivity and subsequently followed up their reproductive performance in the wild. Problem-solver females produced larger clutches than nonsolvers. This benefit did not arise because solvers timed their breeding better, occupied better habitats, or compromised offspring quality or their own survival. Instead, foraging range size and day length were relatively small and short, respectively, for solvers, suggesting that they were more efficient at exploiting their environment. In contrast to the positive effect on clutch size, problem solvers deserted their nests more often, leading to little or no overall selection on problem-solving performance. Our results are consistent with the idea that variation in cognitive ability is shaped by contrasting effects on different life history traits directly linked to fitness [1, 3].