Long-term memory in wild falcons.

Long-term memory - information retention over long timescales - can allow animals to retain foraging skills and efficiently respond to seasonally available resources and changing environments1. Most long-term memory research is with captive species, focusing on spatial, individual or object recognition, with less known about wild species and the retention of motor task abilities, as in the case of complex foraging skills2,3. We have examined whether wild striated caracaras (Phalcoboenus australis), recently shown to rapidly and flexibly innovate with an eight-task puzzle box4, retain task memories one year later. We found that, despite no reinforcement, caracaras repeated motor techniques that led to their most recent success on tasks the year prior, solving nearly twice as fast as a naïve control group and four times faster than when naïve. Our results suggest long-term memory may be important for non-migratory opportunistic generalists, particularly in remote island environments with seasonally available resources, and further highlight how striated caracaras are promising candidates for avian cognitive studies.

Putting the best foot forward: Limb lateralization in the Goffin's cockatoo (Cacatua goffiniana).

Many parrot species exhibit a high degree of limb lateralization on both the individual and species levels. In particular, the members of the cockatoo family are left-footed for food-holding at proportions reminiscent of right-handedness in humans. Here, we examine the limb lateralization of the Goffin's cockatoo (Cacatua goffiniana), a tool-using and technically proficient species used as a model of parrot cognition. First, we investigated the postural origins theory, originally proposed in primates to explain handedness. According to this theory, the hand that was used by ancestral primates to cling to trees developed finer motor control. Using a series of problem-solving tasks, we tested the possibility that the parrot's postural foot, which is similarly used to grasp tree branches, could be more motorically skilled. Although we did not find support for this idea, we did discover that task type does affect foot use, as subjects switched from using their food-holding dominant foot to their other foot during reaching tasks. We also found that the cockatoos more flexibly switched and used both feet when faced with more challenging tasks. Secondly, we attempted a partial replication of a previous study with parrots derived from the enhanced cognition hypothesis, which claimed that more lateralized individuals were better problem solvers. However, we did not find this relationship to be significant in any of our tasks. We did confirm that individual Goffin's cockatoos are extremely limb lateralized for food-holding in addition to other tasks, which may play a role in their approaches to problem-solving. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Children's limited tooling ability in a novel concurrent tool use task supports the innovation gap.

School-aged children have consistently shown a surprising developmental lag when attempting to innovate solutions to tool use tasks, despite being capable of learning to solve these problems from a demonstrator. We suggest that this "innovation gap" arises from tool tasks with more complex spatial relations. Following Fragaszy and Mangalam's new tooling theory, we predicted that innovating a new "sticker slide" task should be more challenging when two tools need to be used at the same time (concurrently) rather than one at a time (sequentially), despite the similarity of the other task elements. In line with previous work, both versions of the task were challenging for all ages of children (4-9 years) that we tested. However, the youngest group showed particularly extreme difficulties, which was marked by not a single child innovating the concurrent version. Although success significantly increased with age, even the oldest group failed to reach 50% success on the concurrent version of the task, whereas the majority of the two older groups could solve the sequential version. Thus, in this first study of concurrent tool use in children, we found support for the prediction that increasing the complexity of spatial relations in tooling exacerbates the innovation gap.

The effect of four different object properties on latency to approach in Goffin's cockatoos (Cacatua goffiniana).

Neophobia and neophilia can be lifesaving as they can facilitate foraging while avoiding predation or intoxication. We investigated the extent to which Goffin's cockatoos (Cacatua goffiniana) exhibit ecollogically relevant and quantifiable neophobic responses toward specific object properties. Twelve cockatoos were presented with 12 novel objects grouped into four distinct categories with unique features: size, color, reflective capacity, and shape. The cockatoos were tested by measuring their latency to approach a high-quality food reward for both novel and control scenarios. Age and sex did not affect the latency to approach food in the presence of a novel object in this species. Additionally, we found no significant differences between the objects of the reflective and color categories. This result is likely due to the plasticity of neophobic behavior related to the benefits and costs of approaching novel stimuli. The cockatoos were significantly slower to approach food in the presence of objects larger than their body size than objects of a similar or smaller size, a phenomenon possibly explained by the increased risk of approaching unknown objects large enough to be a potential predator. They were also significantly more hesitant to approach food in the presence of elongated objects, a phenomenon potentially explained by an ecologically relevant avoidance of snakes. The extent of this neophobia was statistically similar at a group level, indicating that avoidance of elongated and large objects could be an adaptive response aiding survival under natural circumstances and that snakes may impose strong selective pressures on this species. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

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.

Ratcheting up tool innovation in Goffin's cockatoos (Cacatua goffiniana): The effect of contextually diverse prior experience.

The ability to gain information from one situation, acquire new skills and/or perfect existing ones, and subsequently apply them to a new situation is a key element in behavioural flexibility and a hallmark of innovation. A flexible agent is expected to store these skills and apply them to contexts different from that in which learning occurred. Goffin's cockatoos (Cacatua goffiniana) are highly innovative parrots renowned for their problem-solving and tool-using skills and are thus excellent candidates to study this phenomenon. We hypothesized that birds allowed to use a tool in a larger variety of contingencies would acquire a broader expertise in handling it, facilitating its transfer to new tasks. In our study, we compared the performance of two groups of captive Goffin's cockatoos (N = 13): A test group received more diverse learning and motor experiences on multiple applications of a hook-type tool, while a control group received intensive, total trial-matched, experience with a single application of the same tool. Then, both groups were tested on two novel tasks to determine whether experience with the tool in multiple contexts would facilitate performance during transfer. While both groups transferred to both novel tasks, group differences in performance were apparent, particularly in the second transfer task, where test birds achieved a higher success rate and reached criteria within fewer trials than control birds. These results provide support for the prediction that experiencing a diverse range of contingencies with a tool appears to allow birds to acquire generalizable knowledge and transferrable skills to tackle an untrained problem more efficiently. In contrast, intensive experience with the tool in a single context might have made control birds less flexible and more fixated on previously learned tool-dependent instances.

Flexible tool set transport in Goffin's cockatoos.

The use of tool sets constitutes one of the most elaborate examples of animal technology, and reports of it in nature are limited to chimpanzees and Goffin's cockatoos. Although tool set use in Goffin's was only recently discovered, we know that chimpanzees flexibly transport tool sets, depending on their need. Flexible tool set transport can be considered full evidence for identification of a genuine tool set, as the selection of the second tool is not just a response to the outcomes of the use of the first tool but implies recognizing the need for both tools before using any of them (thus, categorizing both tools together as a tool set). In three controlled experiments, we tested captive Goffin's in tasks inspired by the termite fishing of Goualougo Triangle's chimpanzees. Thereby, we show that some Goffin's can innovate the use and flexibly use and transport a new tool set for immediate future use; therefore, their sequential tool use is more than the sum of its parts. VIDEO ABSTRACT.

The Innovation Arena: A Method for Comparing Innovative Problem-Solving Across Groups.

Problem-solving tasks are commonly used to investigate technical, innovative behavior but a comparison of this ability across a broad range of species is a challenging undertaking. Specific predispositions, such as the morphological toolkit of a species or exploration techniques, can substantially influence performance in such tasks, which makes direct comparisons difficult. The method presented here was developed to be more robust with regard to such species-specific differences: the Innovation Arena presents 20 different problem-solving tasks. All tasks are presented simultaneously. Subjects are confronted with the apparatus repeatedly, which allows a measurement of the emergence of innovations over time - an important next step for investigating how animals can adapt to changing environmental conditions through innovative behavior. Each individual was tested with the apparatus until it ceased to discover solutions. After testing was concluded, we analyzed the video recordings and coded successful retrieval of rewards and multiple apparatus-directed behaviors. The latter were analyzed using a Principal Component Analysis and the resulting components were then included in a Generalized Linear Mixed Model together with session number and the group comparison of interest to predict the probability of success. We used this approach in a first study to target the question of whether long-term captivity influences the problem-solving ability of a parrot species known for its innovative behavior: the Goffin´s cockatoo. We found an effect in degree of motivation but no difference in the problem-solving ability between short- and long-term captive groups.

Innovative composite tool use by Goffin's cockatoos (Cacatua goffiniana).

Composite tool use (using more than one tool simultaneously to achieve an end) has played a significant role in the development of human technology. Typically, it depends on a number of specific and often complex spatial relations and there are thus very few reported cases in non-human animals (e.g., specific nut-cracking techniques in chimpanzees and capuchin monkeys). The innovative strategies underlying the innovation and spread of tool manufacture and associative tool use (using > 1 tools) across tool using animals is an important milestone towards a better understanding of the evolution of human technology. We tested Goffin's cockatoos on a composite tool problem, the 'Golf Club Task', that requires the use of two objects in combination (one used to control the free movement of a second) to get a reward. We demonstrate that these parrots can innovate composite tool use by actively controlling the position of the end effector and movement of both objects involved in a goal directed manner. The consistent use of different techniques by different subjects highlights the innovative nature of the individual solutions. To test whether the solution could be socially transmitted, we conducted a second study, which provided only tentative evidence for emulative learning. To our knowledge, this indicates that the cognitive preconditions for composite tool use have also evolved outside the primate lineage.

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.

Wild Goffin's cockatoos flexibly manufacture and use tool sets.

The use of different tools to achieve a single goal is considered unique to human and primate technology. To unravel the origins of such complex behaviors, it is crucial to investigate tool use that is not necessary for a species' survival. These cases can be assumed to have emerged innovatively and be applied flexibly, thus emphasizing creativity and intelligence. However, it is intrinsically challenging to record tool innovations in natural settings that do not occur species-wide. Here, we report the discovery of two distinct tool manufacture methods and the use of tool sets in wild Goffin's cockatoos (Cacatua goffiniana). Up to three types of wooden tools, differing in their physical properties and each serving a different function, were manufactured and employed to extract embedded seed matter of Cerbera manghas. While Goffin's cockatoos do not depend on tool-obtained resources, repeated observations of two temporarily captive wild birds and indications from free-ranging individuals suggest this behavior occurs in the wild, albeit not species-wide. The use of a tool set in a non-primate implies convergent evolution of advanced tool use. Furthermore, these observations demonstrate how a species without hands can achieve dexterity in a high-precision task. The presence of flexible use and manufacture of tool sets in animals distantly related to humans significantly diversifies the phylogenetic landscape of technology and opens multiple avenues for future research. VIDEO ABSTRACT.

Goffin's Cockatoos (Cacatua goffiniana) Can Solve a Novel Problem After Conflicting Past Experiences.

Novel problems often partially overlap with familiar ones. Some features match the qualities of previous situations stored in long-term memory and therefore trigger their retrieval. Using relevant, while inhibiting irrelevant, memories to solve novel problems is a hallmark of behavioral flexibility in humans and has recently been demonstrated in great apes. This capacity has been proposed to promote technical innovativeness and thus warrants investigations of such a mechanism in other innovative species. Here, we show that proficient tool-users among Goffin's cockatoos-an innovative tool-using species-could use a relevant previous experience to solve a novel, partially overlapping problem, even despite a conflicting, potentially misleading, experience. This suggests that selecting relevant experiences over irrelevant experiences guides problem solving at least in some Goffin's cockatoos. Our result supports the hypothesis that flexible memory functions may promote technical innovations.

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.

String-pulling in the Goffin's cockatoo (Cacatua goffiniana).

Goffin's cockatoos, a parrot species endemic to the Tanimbar Islands in Indonesia, demonstrate remarkable cognitive skills across various technical tasks. These neophilic extractive foragers explore objects with their beak and feet, and are skilled in several modes of tool use. In this study, we confronted the animals for the first time with a vertical string-pulling setup, including a set of classic and novel controls. Nine of the 12 subjects, two of which were subadults, immediately interacted with the single-string task, with seven individuals successfully obtaining the reward on their very first attempt. Four different double string discrimination tests with varying spatial relations were used to assess the Goffin's cockatoos' apprehension of basic physical task properties. We found significant differences in performance between the respective experimental conditions, as well as the development of side biases. The results suggest that while the birds seem to consider simple cause-effect relationships, there is no evidence for a mental representation of the causal mechanisms underlying the string-pulling tasks, as subjects failed the crossed strings condition out of immediate sight. Finally, we provide suggestions on improving the methodology, and discuss our findings in regard to the Goffin's cockatoo's ecology.

Current Understanding of the "Insight" Phenomenon Across Disciplines.

Despite countless anecdotes and the historical significance of insight as a problem solving mechanism, its nature has long remained elusive. The conscious experience of insight is notoriously difficult to trace in non-verbal animals. Although studying insight has presented a significant challenge even to neurobiology and psychology, human neuroimaging studies have cleared the theoretical landscape, as they have begun to reveal the underlying mechanisms. The study of insight in non-human animals has, in contrast, remained limited to innovative adjustments to experimental designs within the classical approach of judging cognitive processes in animals, based on task performance. This leaves no apparent possibility of ending debates from different interpretations emerging from conflicting schools of thought. We believe that comparative cognition has thus much to gain by embracing advances from neuroscience and human cognitive psychology. We will review literature on insight (mainly human) and discuss the consequences of these findings to comparative cognition.

Using an Innovation Arena to compare wild-caught and laboratory Goffin's cockatoos.

The ability to innovate, i.e., to exhibit new or modified learned behaviours, can facilitate adaptation to environmental changes or exploiting novel resources. We hereby introduce a comparative approach for studying innovation rate, the 'Innovation Arena' (IA), featuring the simultaneous presentation of 20 interchangeable tasks, which subjects encounter repeatedly. The new design allows for the experimental study of innovation per time unit and for uncovering group-specific problem-solving abilities - an important feature for comparing animals with different predispositions and life histories. We applied the IA for the first time to investigate how long-term captivity affects innovative capacities in the Goffin's cockatoo, an avian model species for animal innovation. We found that fewer temporarily-captive wild birds are inclined to consistently interact with the apparatus in comparison to laboratory-raised birds. However, those that are interested solve a similar number of tasks at a similar rate, indicating no difference in the cognitive ability to solve technical problems. Our findings thus provide a contrast to previous literature, which suggested enhanced cognitive abilities and technical problem-solving skills in long-term captive animals. We discuss the impact and discrepancy between motivation and cognitive ability on innovation rate. Our findings contribute to the debate on how captivity affects innovation in animals.

Orangutans (Pongo abelii) make flexible decisions relative to reward quality and tool functionality in a multi-dimensional tool-use task.

Making economic decisions in a natural foraging situation that involves the use of tools may require an animal to consider more levels of relational complexity than merely deciding between an immediate and a delayed food option. We used the same method previously used with Goffin´s cockatoos to investigate the orangutans' flexibility for making the most profitable decisions when confronted with five different settings that included one or two different apparatuses, two different tools and two food items (one more preferred than the other). We found that orangutans made profitable decisions relative to reward quality, when the task required the subjects to select a tool over an immediately accessible food reward. Furthermore, most subjects were sensitive to work-effort when the immediate and the delayed option (directly accessible by using a tool) led to the same outcome. Most subjects continued to make profitable decisions that required taking into account the tool functionality. In a final multidimensional task design in which subjects had to simultaneously focus on two apparatuses, two reward qualities and two different tools, the orangutans chose the functional tool to access the high quality reward.

Goffin's cockatoos make the same tool type from different materials.

Innovative tool manufacture is rare and hard to isolate in animals. We show that an Indonesian generalist parrot, the Goffin's cockatoo, can flexibly and spontaneously transfer the manufacture of stick-type tools across three different materials. Each material required different manipulation patterns, including substrates that required active sculpting for achieving a functional, elongated shape.

Inference by Exclusion in Goffin Cockatoos (Cacatua goffini).

Inference by exclusion, the ability to base choices on the systematic exclusion of alternatives, has been studied in many nonhuman species over the past decade. However, the majority of methodologies employed so far are hard to integrate into a comparative framework as they rarely use controls for the effect of neophilia. Here, we present an improved approach that takes neophilia into account, using an abstract two-choice task on a touch screen, which is equally feasible for a large variety of species. To test this approach we chose Goffin cockatoos (Cacatua goffini), a highly explorative Indonesian parrot species, which have recently been reported to have sophisticated cognitive skills in the technical domain. Our results indicate that Goffin cockatoos are able to solve such abstract two-choice tasks employing inference by exclusion but also highlight the importance of other response strategies.