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Adaptive theory predicts that the fundamental trade-off between starvation and predation risk shapes diurnal patterns in foraging activity and mass gain in wintering passerine birds. Foragers mitigating both types of risk should exhibit a bimodal distribution (increased foraging and mass gain early and late in the day), whereas both foraging and mass gains early (versus late) during the day are expected when the risk of starvation (versus predation) is greatest. Finally, relatively constant rates of foraging and mass gain should occur when the starvation–predation risk trade-off is independent of body mass. Using automated feeders with integrated digital balances, we estimated diurnal patterns in foraging and body mass gain to test which ecological scenario was best supported in wintering great tits Parus major. Based on data of 40 consecutive winter days recording over 12 000 body masses of 28 individuals, we concluded that birds foraged and gained mass early during the day, as predicted by theory when the starvation–predation risk trade-off is mass-dependent and starvation risk outweighs predation risk. Slower explorers visited the feeders more often, and decreased their activity along the day more strongly, compared with faster explorers, thereby explaining a major portion of the individual differences in diurnal patterning of foraging activity detected using random regression analyses. Birds did not differ in body mass gain trajectories, implying both that individuals differed in the usage of feeders, and that unbiased conclusions regarding how birds resolve starvation–predation risk trade-off require the simultaneous recording of foraging activity and body mass gain trajectories. Our study thereby provides the first unambiguous demonstration that individual birds are capable of adjusting their diurnal foraging and mass gain trajectories in response to ecological predictors of starvation risk as predicted by starvation–predation risk trade-off theory.

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.

Zebrafish ( Danio rerio ) are widely used in behavioural neuroscience as a model for studying anxiety-like and stress-related behaviours. However, substantial variability exists within and among individuals, influenced by factors such as sex, age, and environmental conditions, making the interpretation of anxiety-related behaviours challenging. Here we characterized longitudinal patterns of stability and variability in anxiety-like behaviours across individual adult zebrafish and assessed whether distinct behavioural profiles emerged over time. Using the novel tank dive test, we tracked anxiety-related behaviours in zebrafish across multiple time points over a 21-week period (90, 120, and 150 days post-fertilization). Behavioural metrics, including time spent in tank zones, swimming velocity, and immobility, were analyzed for age- and sex-related effects, repeatability, and group variation. Results indicated significant changes in anxiety-like behaviours with age, with fish spending more time in the upper zone and displaying increased swimming velocity over time. While no significant sex differences were observed in zone preference, males exhibited greater within-individual variation in time spent in the lower zone, while females demonstrated higher among-individual variation and repeatability over time. Furthermore, zebrafish were classified into high, medium, and low-anxiety groups based on cumulative behavioural scores, revealing stable individual differences in anxiety-like behaviours. These findings highlight the importance of considering age, sex and both intra- and inter-individual variation when interpreting zebrafish behaviour and provide a foundation for future research exploring selective breeding, anxiety level interactions, and pharmacological modulation of anxiety-related phenotypes.

Despite a wealth of studies documenting prey responses to perceived predation risk, researchers have only recently begun to consider how prey integrate information from multiple cues in their assessment of risk. We conduct a systematic review and meta-analysis of studies that experimentally manipulated perceived predation risk in birds and evaluate support for three alternative models of cue integration: redundancy/equivalence, enhancement, and antagonism. One key insight from our analysis is that the current theory, generally applied to study cue integration in animals, is incomplete. These theories specify the effects of increasing information level on mean, but not variance, in responses. In contrast, we show that providing multiple complementary cues of predation risk simultaneously does not affect mean response. Instead, as information richness increases, populations appear to assess risk more accurately, resulting in lower among-population variance in response to manipulations of perceived predation risk. We show that this may arise via a statistical process called maximum-likelihood estimation (MLE) integration. Our meta-analysis illustrates how explicit consideration of variance in responses can yield important biological insights. Animal behavioural response to predation risk could depend on the type and number of cues. This global metaanalysis shows that providing multiple cues of predation risk reduces variance in the behavioural responses of birds.

Phenotypic plasticity is a major mechanism whereby organisms adjust their traits within‐generations to changes in environmental conditions. In the context of range expansions, plasticity is thought to be especially important, as plastic changes in traits can lead to rapid adaptation. One epigenetic process in particular, DNA methylation, enables organisms to adjust gene expression contingent on the environment, which suggests it may play a role in range expansions. At present, we know little about how methylation is regulated in wildlife, especially expression of the enzymes responsible for altering methyl marks on the genome. In this study, we compared expression of three epigenetic regulator genes (DNA methyltransferase 1, DNMT1; DNA methyltransferase 3, DNMT3; and one ten‐eleven translocation methylcytosine dioxygenase, TET2) in three tissues (gut, liver, and spleen) of house sparrows Passer domesticus from nine countries. Some countries are in the native range of the species (Israel, the Netherlands, Norway, Spain, and Vietnam) whereas others are sites the species has colonized in the last 150 years (i.e. Australia, Canada, New Zealand, and Senegal). In this exploratory study, we asked whether non‐native birds and/or birds from sites with comparatively unpredictable climates would express different levels of these genes. We found that all three genes were expressed more in sparrows from the native range and from areas with more stable temperatures. Expression of all three genes was also strongly correlated among‐locations and within‐individuals, but mean expression was quite different among tissues. Many factors (e.g. urbanization of the capture site, sex of the bird) did not significantly affect gene expression, but others surprisingly did (e.g. latitude). Our results suggest that these enzymes could be important in range expansions or geographic distribution generally, but more detailed investigations will be insightful.

Prey animals must accurately assess predation risk within their environment. To gather information about this risk, prey animals may personally sample the environment (“personal information”) or observe the behavior of congeners (“social information”). Personal information is thought to be more accurate and reliable but may also require more time and energy to acquire. On the other hand, social information, such as alarm calls, tends to be less costly to obtain but may also be less reliable if congeners assess risk differently from one another, or if the information quickly becomes outdated. Theoretical models predict that individuals will differ in how they value personal versus social information. We used previously collected data from a marked population of black‐capped chickadees (Poecile atricapillus) to test this prediction. Chickadees were exposed to three different predator cue types: a predator mount (personal information), conspecific mobbing calls (social information), and a combination of both (personal + social information) near feeders established on their territory. We recorded the time it took a chickadee to visit a feeder following cue exposure (i.e., latency to resume feeding) to evaluate individual differences in response to predator cues. Contrary to our prediction, we found no evidence that individuals differed in how they valued personal versus social information about predation risk. Instead, our results suggest that responses to predator cues are state‐dependent, with some individuals consistently responding more strongly than others, regardless of cue type. We also found that when chickadees were exposed to a combination of social and personal predator cues, they exhibited higher among‐individual variation in latency to resume feeding than when they were exposed to social or personal cues alone. We discuss how individual differences in cue integration (i.e., cue redundancy/equivalence, enhancement, and antagonism) may account for this finding. Chickadees can use different sources of information (e.g., personal vs. social) to evaluate predation risk. Individual responses to different sources of information were positively correlated; birds that responded strongly to personal information also responded strongly to social. Combined cues led to higher variation among individuals, suggesting that individuals differ in how they integrate social versus personal information cues.

Foragers can gain knowledge of profitable foraging opportunities either by sampling the environment directly (asocial information) or from congeners (social information). The relative benefit of using social information over asocial information is context-specific, and social information use is expected to be particularly beneficial when the costs of acquiring asocial information are high, for example, due to high risk of starvation if asocial information fails. We investigated the plasticity of social information use in an overwintering population of black-capped chickadees ( Poecile atricapillus ) as they rediscovered an intermittently available food source. Lower temperatures impose energetic costs that increase the risk of starvation in chickadees; therefore, lower temperatures are predicted to favour higher use of social information. To test this prediction, we evaluated chickadees’ reliance on social information during foraging as ambient temperatures ranged from −11.0°C to 5.5°C. We evaluated the relative strength of reliance on social and asocial information using network-based diffusion analysis. We found increased reliance on social information transmission with decreasing temperature. Reversible plasticity of social information use may be an important mechanism to cope with low ambient temperatures, a seasonal challenge experienced by many animals.

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. We investigated lethal (mortality) and nonlethal (body condition) effects of different two PIT tagging methods (implants or leg band embedded) compared with birds that were color banded but not fitted with PIT tags. In a sample of over 200 birds followed for more than two years, we found no evidence of differences in survival or body condition across any of these three treatments.

Variation in life history (LH) traits along the fast-slow continuum (referred to as pace of life, POL) is thought to result from a trade-off between investments in current versus future reproduction. Originally developed for understanding variation in LH strategies at the among-population level, the POL theory has more recently been applied towards understanding variation in LH traits at the withinpopulation level, and further extended to address the covariance of LH traits with additional behavioural and/or physiological traits, referred to as pace-of-life syndromes (POLS). The article by Reale et al. (Philos T Roy Soc 365:4051-4063, 2010), which synthesized several earlier reviews and opinions on among-individual covariation between LH, behavioural, and physiological traits, and subsequent research testing POLS in a variety of species, have collectively been cited several hundreds of times—a trend that continues. These works have interdisciplinary impact, informing research in life history biology, behavioural and developmental biology, and the social sciences. In this paper, we review the existing theoretical POLS models that provide adaptive explanations for covariances between LH traits and additional behavioural and/or physiological traits while assuming a trade-off between current and future reproduction. We find that the set of relevant models is small. Moreover, models show that covariances between life history traits and behavioural or physiological traits can arise even in the absence of a current-future reproduction trade-off, implying that observing such covariances does not provide a strong indication regarding the process generating POLS. We discuss lessons learned from existing models of POLS, highlight key gaps in the modelling literature, and provide guidelines for better integration between theory and data.

We show that a higher vertebrate can graze surficial intertidal biofilm, previously only considered a food source for rasping invertebrates and a few specialized fish. Using evidence from video recordings, stomach contents, and stable isotopes, we describe for the first time the grazing behavior of Western Sandpipers (Calidris mauri) and estimate that biofilm accounts for 45—59% of their total diet or 50% of their daily energy budget. Our finding of shorebirds as herbivores extends the trophic range of shorebirds to primary consumers and potential competitors with grazing invertebrates. Also, given individual grazing rates estimated at seven times body mass per day and flock sizes into the tens of thousands, biofilm-feeding shorebirds could have major impacts on sediment dynamics. We stress the importance of the physical and biological processes maintaining biofilm to shorebird and intertidal conservation.