Animal culture research should include avian nest construction.

Material culture-that is, group-shared and socially learned object-related behaviour(s)-is a widespread and diverse phenomenon in humans. For decades, researchers have sought to confirm the existence of material culture in non-human animals; however, the main study systems of interest-namely, tool making and/or using non-human primates and corvids-cannot provide such confirmatory evidence: because long-standing ethical and logistical constraints handicap the collection of necessary experimental data. Synthesizing evidence across decades and disciplines, here, I present a novel framework for (mechanistic, developmental, behavioural, and comparative) study on animal material culture: avian nest construction.

Social learning about construction behaviour via an artefact.

One source of public information may be the enduring products of others' behaviour, such as discarded tools or vacated nests. Here, we examined whether observation of a nest affects the material captive zebra finch males prefer when they construct their first nest. It does: for first-time nest construction, males that viewed only an empty cage preferred the colour of material each initially favoured but those males that had observed a pre-built nest of material of their non-preferred colour lost their material-colour preference altogether. Additionally, half of the males that viewed a nest were tested in an environment (the laboratory) different to that in which they were reared (an outdoor aviary). We had expected the aviary-reared (versus laboratory-reared) males would be more uncertain, and thus more likely to select material for their first nest that matched in colour to the colour of the 'demonstrated' nest-but this was not the case. The aviary-reared males did, however, tend to touch first the demonstrated colour of material more than did the laboratory-reared males. Together these results show that both observation of a nest and a change in environment can influence the material choices of novice builders. For naïve animal builders, then, construction artefacts can be information resources for learning about potential construction material.

How New Caledonian crows solve novel foraging problems and what it means for cumulative culture.

New Caledonian crows make and use tools, and tool types vary over geographic landscapes. Social learning may explain the variation in tool design, but it is unknown to what degree social learning accounts for the maintenance of these designs. Indeed, little is known about the mechanisms these crows use to obtain information from others, despite the question's importance in understanding whether tool behavior is transmitted via social, genetic, or environmental means. For social transmission to account for tool-type variation, copying must utilize a mechanism that is action specific (e.g., pushing left vs. right) as well as context specific (e.g., pushing a particular object vs. any object). To determine whether crows can copy a demonstrator's actions as well as the contexts in which they occur, we conducted a diffusion experiment using a novel foraging task. We used a nontool task to eliminate any confounds introduced by individual differences in their prior tool experience. Two groups had demonstrators (trained in isolation on different options of a four-option task, including a two-action option) and one group did not. We found that crows socially learn about context: After observers see a demonstrator interact with the task, they are more likely to interact with the same parts of the task. In contrast, observers did not copy the demonstrator's specific actions. Our results suggest it is unlikely that observing tool-making behavior transmits tool types. We suggest it is possible that tool types are transmitted when crows copy the physical form of the tools they encounter.

Risk-sensitive learning is a winning strategy for leading an urban invasion.

In the unpredictable Anthropocene, a particularly pressing open question is how certain species invade urban environments. Sex-biased dispersal and learning arguably influence movement ecology, but their joint influence remains unexplored empirically, and might vary by space and time. We assayed reinforcement learning in wild-caught, temporarily captive core-, middle-, or edge-range great-tailed grackles-a bird species undergoing urban-tracking rapid range expansion, led by dispersing males. We show, across populations, both sexes initially perform similarly when learning stimulus-reward pairings, but, when reward contingencies reverse, male-versus female-grackles finish 'relearning' faster, making fewer choice-option switches. How do male grackles do this? Bayesian cognitive modelling revealed male grackles' choice behaviour is governed more strongly by the 'weight' of relative differences in recent foraging payoffs-i.e., they show more pronounced risk-sensitive learning. Confirming this mechanism, agent-based forward simulations of reinforcement learning-where we simulate 'birds' based on empirical estimates of our grackles' reinforcement learning-replicate our sex-difference behavioural data. Finally, evolutionary modelling revealed natural selection should favour risk-sensitive learning in hypothesised urban-like environments: stable but stochastic settings. Together, these results imply risk-sensitive learning is a winning strategy for urban-invasion leaders, underscoring the potential for life history and cognition to shape invasion success in human-modified environments.

Reproductive consequences of material use in avian nest construction.

Birds' nests represent a rich behavioural 'fingerprint', comprising several important decisions-not the least of which is the selection of appropriate material. Material selection in nest-building birds is thought to reflect, in part, builder-birds' use of the 'best' material-in terms of physical properties (e.g., rigidity)-refined across generations. There is, however, little experimental evidence to link the physical properties of nest material to both birds' nest-building and breeding performance. We examined individual-level material-use consequences for breeding zebra finches by manipulating the kind of material available to laboratory-housed pairs: stiff or flexible same-length string. We show that higher fledgling numbers were related to: (i) fewer pieces used in nest construction by stiff-string builders; and conversely, (ii) more pieces used in nest construction by flexible-string builders. Together, these data suggest that physical differences in nest material can affect avian reproduction (here, the trade-off between nest-construction investment and fledgling success), highlighting the adaptive significance of nest-building birds' material selectivity.

Modifications to the Aesop's Fable paradigm change New Caledonian crow performances.

While humans are able to understand much about causality, it is unclear to what extent non-human animals can do the same. The Aesop's Fable paradigm requires an animal to drop stones into a water-filled tube to bring a floating food reward within reach. Rook, Eurasian jay, and New Caledonian crow performances are similar to those of children under seven years of age when solving this task. However, we know very little about the cognition underpinning these birds' performances. Here, we address several limitations of previous Aesop's Fable studies to gain insight into the causal cognition of New Caledonian crows. Our results provide the first evidence that any non-human animal can solve the U-tube task and can discriminate between water-filled tubes of different volumes. However, our results do not provide support for the hypothesis that these crows can infer the presence of a hidden causal mechanism. They also call into question previous object-discrimination performances. The methodologies outlined here should allow for more powerful comparisons between humans and other animal species and thus help us to determine which aspects of causal cognition are distinct to humans.