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.

Individual risk-taking behaviour in black-capped chickadees (Poecile atricapillus) does not predict annual survival.

Within species, individuals often show repeatable differences in behaviours, called 'animal personality'. One behaviour that has been widely studied is how quickly an individual resumes feeding after a disturbance, referred to as boldness or risk-taking. Depending on the mechanism(s) shaping risk-taking behaviour, risk-taking could be positively, negatively, or not associated with differences in overall survival. We studied risk-taking and survival in a population of free-living black-capped chickadees (Poecile atricapillus) in which we previously showed repeatable among-individual differences in risk-taking over the course of several months. We found no evidence that variation in risk-taking is associated with differences in annual survival. We suggest that variation in risk-taking is likely shaped by multiple mechanisms simultaneously, such that the net effect on survival is small or null. For example, among-individual differences in energy demand may favour greater risk-taking without imposing an overall mortality cost if higher energy demand covaries with escape flight performance. We propose directions for future work, including using a multi-trait, multi-year approach to study risk-taking, to allow for stronger inferences regarding the mechanisms shaping these behavioural decisions.

Visual cues of predation risk outweigh acoustic cues: a field experiment in black-capped chickadees.

Foragers rely on various cues to assess predation risk. Information theory predicts that high certainty cues should be valued more than low certainty cues. We measured the latency of black-capped chickadees (Poecile atricapillus) to resume feeding during winter in response to cues that conferred different degrees of certainty about current predation risk: a high certainty visual cue (predator mount) and a lower certainty acoustic cue (conspecific mobbing calls), presented either alone or in combination. As predicted, chickadees took longer to resume feeding after the visual than the acoustic cue, and this effect was greatest under conditions of high starvation risk (i.e. low temperatures). Presenting both cues together produced the same foraging delay as the visual cue alone under low starvation risk, but surprisingly, resulted in lower responses under high starvation risk compared to the visual cue alone. We suggest that this may be due to prey using a form of information updating, whereby differences in the timing of perception of acoustic versus visual cues interacts with energetic constraint to shape perceived risk. Although the sequential perception of cues is likely in a range of decision-making contexts, studies manipulating the order in which cues are perceived are needed to test existing models of multimodal cue integration.

Effects of manipulated levels of predation threat on parental provisioning and nestling begging.

Parental provisioning behavior is a major determinant of offspring growth and survival, but high provisioning rates might come at the cost of increased predation threat. Parents should thus adjust provisioning activity according to current predation threat levels. Moreover, life-history theory predicts that response to predation threat should be correlated with investment in current reproduction. We experimentally manipulated perceived predation threat in free-living great tits (Parus major) by presenting parents with a nest predator model while monitoring different aspects of provisioning behavior and nestling begging. Experiments were conducted in 2 years differing greatly in ecological conditions, including food availability. We further quantified male territorial aggressiveness and male and female exploratory tendency. Parents adjusted provisioning according to current levels of threat in an apparently adaptive way. They delayed nest visits during periods of elevated perceived predation threat and subsequently compensated for lost feeding opportunities by increasing provisioning once the immediate threat had diminished. Nestling begging increased after elevated levels of predation threat, but returned to baseline levels by the end of the experiment, suggesting that parents had fully compensated for lost feeding opportunities. There was no evidence for a link between male exploration behavior or aggressiveness and provisioning behavior. In contrast, fast-exploring females provisioned at higher rates, but only in the year with poor environmental conditions, which might indicate a greater willingness to invest in current reproduction in general. Future work should assess whether these personality-related differences in delivery rates under harsher conditions came at a cost of reduced residual reproductive value.

Sources of (co)variation in alternative siring routes available to male great tits (Parus major).

Males of socially monogamous species can increase their siring success via within-pair and extra-pair fertilizations. In this study, we focused on the different sources of (co)variation between these siring routes, and asked how each contributes to total siring success. We quantified the fertilization routes to siring success, as well as behaviors that have been hypothesized to affect siring success, over a five-year period for a wild population of great tits Parus major. We considered siring success and its fertilization routes as "interactive phenotypes" arising from phenotypic contributions of both members of the social pair. We show that siring success is strongly affected by the fecundity of the social (female) partner. We also demonstrate that a strong positive correlation between extra-pair fertilization success and paternity loss likely constrains the evolution of these two routes. Moreover, we show that more explorative and aggressive males had less extra-pair fertilizations, whereas more explorative females laid larger clutches. This study thus demonstrates that (co)variation in siring routes is caused by multiple factors not necessarily related to characteristics of males. We thereby highlight the importance of acknowledging the multilevel structure of male fertilization routes when studying the evolution of male mating strategies.

Does coping style predict optimization? An experimental test in a wild passerine bird.

A number of studies have suggested that avian brood size is individually optimized. Yet, optimal reproductive decisions likely vary owing to among-individual differences in environmental sensitivity. Specifically, 'proactive' individuals who do not track environmental changes may be less able to produce optimal brood sizes than 'reactive' individuals who have more precise local environmental knowledge. To test this, we quantified exploratory behaviour (a proxy for proactivity) in a great tit (Parus major) population, manipulated brood sizes (reduced, control, enlarged) and evaluated whether individuals of dissimilar coping style differed in their level of optimization. If reactive females behaved optimally, any deviation from their original brood size should lower fitness, whereas this should not be the case for proactive females. Reactive females indeed performed best at their natural brood size, whereas proactive females performed best when raising an enlarged brood. These findings imply that proactive females produced sub-optimal brood sizes. We speculate that proactive females might (i) take decisions based on biased perception of their environment, (ii) face energetic constraints in offspring production and/or (iii) be more willing to invest into current reproduction when given the option. Our findings provide experimental evidence for coping style-related differences in optimal reproductive decisions and life-history strategies.