Information Gathering Is Associated with Increased Survival: A Field Experiment in Black-Capped Chickadees (Poecile atricapillus).

AbstractSampling, investing time or energy to learn about the environment, allows organisms to track changes in resource distribution and quality. The use of sampling is predicted to change as a function of energy expenditure, food availability, and starvation risk, all of which can vary both within and among individuals. We studied sampling behavior in a field study with black-capped chickadees (Poecile atricapillus) and show that individuals adjust their use of sampling as a function of ambient temperature (a proxy for energy expenditure), the presence of an alternative food source (yes or no, a proxy for risk of energy shortfall), and their interaction, as predicted by models of optimal sampling. We also observed repeatable differences in sampling. Some individuals consistently sampled more, and individuals that sampled more overall also had a higher annual survival. These results are consistent with among-individual differences in resource acquisition (e.g., food caches or dominance-related differences in priority access to feeders), shaping among-individual differences in both sampling and survival, with greater resource acquisition leading to both higher sampling and higher survival. Although this explanation requires explicit testing, it is in line with several recent studies suggesting that variation in resource acquisition is a key mechanism underlying animal personality.

Context is key: A comment on Herczeg et al. 2019.

In the last several years, there has been a surge in the number of studies addressing the causes and consequences of among-individual variation in cognitive ability and behavioural plasticity. Here, we use a recent publication by Herczeg et al. (2019: 32(3), 218-226) to highlight three shortcomings common to this newly emerging field. In their study, Herczeg et al. attempted to link variation in cognitive ability and behavioural plasticity by testing whether selection lines of guppies (Poecilia reticulata) that differ in relative brain size also differ in behavioural plasticity, as might be expected if the costs to plasticity are predominantly derived from the cost of developing large brains. First, residual brain size may not be a suitable proxy for 'cognitive ability'. Recent work has shown that intraspecific variation in cognitive ability can be better understood by considering variation in the specific brain areas responsible for the relevant behaviours as opposed to whole-brain mass. Second, the measure of behavioural plasticity, habituation, is unlikely to fulfil the assumptions that plasticity is both adaptive and costly. Finally, we point out several misconceptions regarding animal personality that continue to contribute to the choice of traits that are not well aligned with study objectives. Elucidating the mechanisms underlying among-individual variation in cognition and behavioural plasticity within populations requires integration between behavioural ecology and comparative cognition, and the study system developed by Herczeg et al. has the potential to provide important mechanistic insights. We hope that by articulating and critically appraising the underlying assumptions that are common in these traditionally separate disciplines, a strong foundation can emerge to allow for more fruitful integration of these fields.

Differences in resource acquisition, not allocation, mediate the relationship between behaviour and fitness: a systematic review and meta-analysis.

Within populations, individuals often show repeatable variation in behaviour, called 'animal personality'. In the last few decades, numerous empirical studies have attempted to elucidate the mechanisms maintaining this variation, such as life-history trade-offs. Theory predicts that among-individual variation in behavioural traits could be maintained if traits that are positively associated with reproduction are simultaneously associated with decreased survival, such that different levels of behavioural expression lead to the same net fitness outcome. However, variation in resource acquisition may also be important in mediating the relationship between individual behaviour and fitness components (survival and reproduction). For example, if certain phenotypes (e.g. dominance or aggressiveness) are associated with higher resource acquisition, those individuals may have both higher reproduction and higher survival, relative to others in the population. When individuals differ in their ability to acquire resources, trade-offs are only expected to be observed at the within-individual level (i.e. for a given amount of resource, if an individual increases its allocation to reproduction, it comes at the cost of allocation to survival, and vice versa), while among individuals traits that are associated with increased survival may also be associated with increased reproduction. We performed a systematic review and meta-analysis, asking: (i) do among-individual differences in behaviour reflect among-individual differences in resource acquisition and/or allocation, and (ii) is the relationship between behaviour and fitness affected by the type of behaviour and the testing environment? Our meta-analysis consisted of 759 estimates from 193 studies. Our meta-analysis revealed a positive correlation between pairs of estimates using both survival and reproduction as fitness proxies. That is, for a given study, behaviours that were associated with increased reproduction were also associated with increased survival, suggesting that variation in behaviour at the among-individual level largely reflects differences among individuals in resource acquisition. Furthermore, we found the same positive correlation between pairs of estimates using both survival and reproduction as fitness proxies at the phenotypic level. This is significant because we also demonstrated that these phenotypic correlations primarily reflect within-individual correlations. Thus, even when accounting for among-individual differences in resource acquisition, we did not find evidence of trade-offs at the within-individual level. Overall, the relationship between behaviour and fitness proxies was not statistically different from zero at the among-individual, phenotypic, and within-individual levels; this relationship was not affected by behavioural category nor by the testing condition. Our meta-analysis highlights that variation in resource acquisition may be more important in driving the relationship between behaviour and fitness than previously thought, including at the within-individual level. We suggest that this may come about via heterogeneity in resource availability or age-related effects, with higher resource availability and/or age leading to state-dependent shifts in behaviour that simultaneously increase both survival and reproduction. We emphasize that future studies examining the mechanisms maintaining behavioural variation in populations should test the link between behavioural expression and resource acquisition - both within and among individuals. Such work will allow the field of animal personality to develop specific predictions regarding the mediating effect of resource acquisition on the fitness consequences of individual behaviour.

No effect of passive integrated transponder tagging method on survival or body condition in a northern population of Black-capped Chickadees (Poecile atricapillus).

Passive integrated transponder (PIT) tags allow a range of individual-level data to be collected passively and have become a commonly used technology in many avian studies. Although the potential adverse effects of PIT tags have been evaluated in several species, explicit investigations of their impacts on small (<12 g) birds are limited. This is important, because it is reasonable to expect that smaller birds could be impacted more strongly by application of PIT tags. In this study, we individually marked Black-capped Chickadees (Poecile atricapillus), a small (circa 10 g) passerine, at the University of Alberta Botanic Garden to evaluate potential lethal and sublethal effects of two PIT tagging methods: attachment to leg bands or subcutaneous implantation. We used a Cox proportional hazards model to compare the apparent survival of chickadees with leg band (N = 79) and implanted PIT tags (N = 77) compared with control birds that received no PIT tags (N = 76) over the subsequent 2 years based on mist net recaptures. We used radio-frequency identification (RFID) redetections of leg band PIT tags to evaluate sex-specific survival and increase the accuracy of our survival estimates. We also used a generalized linear regression model to compare the body condition of birds recaptured after overwintering with leg band PIT tags, implanted PIT tags, or neither. Our analysis found no evidence for adverse effects of either PIT tagging method on survival or body condition. While we recommend carefully monitoring study animals and evaluating the efficacy of different PIT tagging methods, we have shown that both leg band and subcutaneously implanted PIT tags ethical means of obtaining individualized information in a small passerine.