Innovative problem solving by wild falcons.

Innovation (i.e., a new solution to a familiar problem, or applying an existing behavior to a novel problem1,2) plays a fundamental role in species' ecology and evolution. It can be a useful measure for cross-group comparisons of behavioral and cognitive flexibility and a proxy for general intelligence.3,4,5 Among birds, experimental studies of innovation (and cognition more generally) are largely from captive corvids and parrots,6,7,8,9,10,11,12 though we lack serious models for avian technical intelligence outside these taxa. Striated caracaras (Phalcoboenus australis) are Falconiformes, sister clade to parrots and passerines,13,14,15 and those endemic to the Falkland Islands (Malvinas) show curiosity and neophilia similar to notoriously neophilic kea parrots16,17 and face similar socio-ecological pressures to corvids and parrots.18,19 We tested wild striated caracaras as a new avian model for technical cognition and innovation using a field-applicable 8-task comparative paradigm (adapted from Rössler et al.20 and Auersperg et al.21). The setup allowed us to assess behavior, rate, and flexibility of problem solving over repeated exposure in a natural setting. Like other generalist species with low neophobia,21,22 we predicted caracaras to demonstrate a haptic approach to solving tasks, flexibly switching to new, unsolved problems and improving their performance over time. Striated caracaras performed comparably to tool-using parrots,20 nearly reaching ceiling levels of innovation in few trials, repeatedly and flexibly solving tasks, and rapidly learning. We attribute our findings to the birds' ecology, including geographic restriction, resource unpredictability, and opportunistic generalism,23,24,25 and encourage future work investigating their cognitive abilities in the wild. VIDEO ABSTRACT.

Socio-ecological correlates of neophobia in corvids.

Behavioral responses to novelty, including fear and subsequent avoidance of novel stimuli, i.e., neophobia, determine how animals interact with their environment. Neophobia aids in navigating risk and impacts on adaptability and survival. There is variation within and between individuals and species; however, lack of large-scale, comparative studies critically limits investigation of the socio-ecological drivers of neophobia. In this study, we tested responses to novel objects and food (alongside familiar food) versus a baseline (familiar food alone) in 10 corvid species (241 subjects) across 10 labs worldwide. There were species differences in the latency to touch familiar food in the novel object and novel food conditions relative to the baseline. Four of seven socio-ecological factors influenced object neophobia: (1) use of urban habitat (versus not), (2) territorial pair versus family group sociality, (3) large versus small maximum flock size, and (4) moderate versus specialized caching (whereas range, hunting live animals, and genus did not), while only maximum flock size influenced food neophobia. We found that, overall, individuals were temporally and contextually repeatable (i.e., consistent) in their novelty responses in all conditions, indicating neophobia is a stable behavioral trait. With this study, we have established a network of corvid researchers, demonstrating potential for further collaboration to explore the evolution of cognition in corvids and other bird species. These novel findings enable us, for the first time in corvids, to identify the socio-ecological correlates of neophobia and grant insight into specific elements that drive higher neophobic responses in this avian family group. VIDEO ABSTRACT.

Goffin's cockatoos discriminate objects based on weight alone.

Paying attention to weight is important when deciding upon an object's efficacy or value in various contexts (e.g. tool use, foraging). Proprioceptive discrimination learning, with objects that differ only in weight, has so far been investigated almost exclusively in primate species. Here, we show that while Goffin's cockatoos learn faster when additional colour cues are used, they can also quickly learn to discriminate between objects on the basis of their weight alone. Ultimately, the birds learned to discriminate between visually identical objects on the basis of weight much faster than primates, although methodological differences between tasks should be considered.

Extending the Reach of Tooling Theory: A Neurocognitive and Phylogenetic Perspective.

Tool use research has suffered from a lack of consistent theoretical frameworks. There is a plethora of tool use definitions and the most widespread ones are so inclusive that the behaviors that fall under them arguably do not have much in common. The situation is aggravated by the prevalence of anecdotes, which have played an undue role in the literature. In order to provide a more rigorous foundation for research and to advance our understanding of the interrelation between tool use and cognition, we suggest the adoption of Fragaszy and Mangalam's (2018) tooling framework, which is characterized by the creation of a body-plus-object system that manages a mechanical interface between tool and surface. Tooling is limited to a narrower suite of behaviors than tool use, which might facilitate its neurocognitive investigation. Indeed, evidence in the literature indicates that tooling has distinct neurocognitive underpinnings not shared by other activities typically classified as tool use, at least in primates. In order to understand the extent of tooling incidences in previous research, we systematically surveyed the comprehensive tool use catalog by Shumaker et al. (2011). We identified 201 tool use submodes, of which only 81 could be classified as tooling, and the majority of the tool use examples across species were poorly supported by evidence. Furthermore, tooling appears to be phylogenetically less widespread than tool use, with the greatest variability found in the primate order. However, in order to confirm these findings and to understand the evolution and neurocognitive mechanisms of tooling, more systematic research will be required in the future, particularly with currently underrepresented taxa.

Investigating information seeking in ravens (Corvus corax).

Measuring the responses of non-human animals to situations of uncertainty is thought to shed light on an animal's metacognitive processes; namely, whether they monitor their own knowledge states. For example, when presented with a foraging task, great apes and macaques selectively seek information about the location of a food item when they have not seen where it was hidden, compared to when they have. We presented this same information seeking task to ravens, in which a food item was hidden in one of three containers, and subjects could either watch where the food was hidden, infer its location through visual or auditory clues, or were given no information. We found that unlike several ape species and macaques, but similar to capuchin monkeys, the ravens looked inside at least one tube on every trial, but typically only once, inside the baited tube, when they had either witnessed it being baited or could visually infer the reward's location. In contrast, subjects looked more often within trials in which they had not witnessed the baiting or were provided with auditory cues about the reward's location. Several potential explanations for these ceiling levels of looking are discussed, including how it may relate to the uncertainty faced by ravens when retrieving food caches.

Do Nonhumans Seek Explanations?

From an early age, children explore their environment in a way suggesting that they reason about causal variables and seek causal explanations. Indeed, following extensive studies of problem-solving abilities in chimpanzees, Povinelli (Folk Physics for Apes, Oxford University Press, 2000) proposed that this ability to reason about unobservable variables is unique to humans. Following on from this, Povinelli and Dunphy-Lelii (Canadian Journal of Experimental Psychology, 55(2), 187-195, 2001) addressed the question whether chimpanzees would explore objects with the aim of elucidating unobservable and surprising object properties. Chimpanzees, unlike preschool children, did not show increased object exploration following a change in the unobservable properties of an object. We critically discuss these findings and argue that more research using a greater variety of methods and with a larger number of species is required to support the hypothesis that only humans engage in explanation seeking. We conclude by highlighting avenues for future research based on developmental and comparative research aimed at object exploration and information seeking conducted since the original investigation by Povinelli and Dunphy-Lelii.

Comparing chimpanzees' preparatory responses to known and unknown future outcomes.

When humans plan for the future, we recognize not only that one particular event may occur, but that the future can have different, mutually exclusive possible outcomes. A recent study by Suddendorf et al (Suddendorf 2017 Biol. Lett.13, 20170170 (doi:10.1098/rsbl.2017.0170)) suggests that young children (less than 3 years) and apes lack this capacity, as demonstrated by their failure to cover each of two tube openings to ensure catching an object that would drop randomly from one of the tubes. Before drawing conclusions based on these negative results, however, it is important to assess subjects' failures and test the reliability of the task itself. To explore whether the apes' performance resulted from an inability to represent mutually exclusive futures or from extraneous factors related to the task, we replicated the methods of Suddendorf et al (Suddendorf 2017 Biol. Lett.13, 20170170 (doi:10.1098/rsbl.2017.0170)) with a group of six chimpanzees but included a key control condition in which subjects were expected to cover both tubes on every trial (i.e. the rewards would consistently emerge from both tubes). We show that even in this straightforward condition in which the outcome of the trial was known, only four of the six subjects ever covered both tubes, typically doing so after a minimum of 17 trials, and often reverting back to covering one tube on later trials. We conclude that this task is not valid for testing the ability to represent mutually exclusive futures. We discuss what potential factors may explain the results and outline a new suggested method to continue testing for this capacity in the future.

Function and flexibility of object exploration in kea and New Caledonian crows.

A range of non-human animals frequently manipulate and explore objects in their environment, which may enable them to learn about physical properties and potentially form more abstract concepts of properties such as weight and rigidity. Whether animals can apply the information learned during their exploration to solve novel problems, however, and whether they actually change their exploratory behaviour to seek functional information about objects have not been fully explored. We allowed kea (Nestor notabilis) and New Caledonian crows (Corvus moneduloides) to explore sets of novel objects both before and after encountering a task in which some of the objects could function as tools. Following this, subjects were given test trials in which they could choose among the objects they had explored to solve a tool-use task. Several individuals from both species performed above chance on these test trials, and only did so after exploring the objects, compared with a control experiment with no prior exploration phase. These results suggest that selection of functional tools may be guided by information acquired during exploration. Neither kea nor crows changed the duration or quality of their exploration after learning that the objects had a functional relevance, suggesting that birds do not adjust their behaviour to explicitly seek this information.

Diffusion of novel foraging behaviour in Amazon parrots through social learning.

While social learning has been demonstrated in species across many taxa, the role it plays in everyday foraging decisions is not well understood. Investigating social learning during foraging could shed light on the emergence of cultural variation in different groups. We used an open diffusion experiment to examine the spread of a novel foraging technique in captive Amazon parrots. Three groups were tested using a two-action foraging box, including experimental groups exposed to demonstrators using different techniques and control birds. We also examined the influence of agonistic and pilfering behaviour on task acquisition. We found evidence of social learning: more experimental birds than control birds interacted with and opened the box. The birds were, however, no more likely to use the demonstrated technique than the non-demonstrated one, making local or stimulus enhancement the most likely mechanism. Exhibiting aggression was positively correlated with box opening, whilst receiving aggression did not reduce motivation to engage with the box, indicating that willingness to defend access to the box was important in task acquisition. Pilfering food and success in opening the box were also positively correlated; however, having food pilfered did not affect victims' motivation to interact with the box. In a group context, pilfering may promote learning of new foraging opportunities. Although previous studies have demonstrated that psittacines are capable of imitation, in this naturalistic set-up there was no evidence that parrots copied the demonstrated opening technique. Foraging behaviour in wild populations of Amazons could therefore be facilitated by low-fidelity social learning mechanisms.

A novel form of spontaneous tool use displayed by several captive greater vasa parrots (Coracopsis vasa).

Parrots are frequently cited for their sophisticated problem-solving abilities, but cases of habitual tool use among psittacines are scarce. We report the first evidence, to our knowledge, of tool use by greater vasa parrots (Coracopsis vasa). Several members of a captive population spontaneously adopted a novel tool-using technique by using pebbles and date pits either (i) to scrape on the inner surface of seashells, subsequently licking the resulting calcium powder from the tool, or (ii) as a wedge to break off smaller pieces of the shell for ingestion. Tool use occurred most frequently just prior to the breeding season, during which time numerous instances of tool transfer were also documented. These observations provide new insights into the tool-using capabilities of parrots and highlight the greater vasa parrot as a species of interest for studies of physical cognition.