Using repeatability of performance within and across contexts to validate measures of behavioral flexibility.

Research into animal cognitive abilities is increasing quickly and often uses methods where behavioral performance on a task is assumed to represent variation in the underlying cognitive trait. However, because these methods rely on behavioral responses as a proxy for cognitive ability, it is important to validate that the task structure does, in fact, target the cognitive trait of interest rather than non-target cognitive, personality, or motivational traits (construct validity). Although it can be difficult, or impossible, to definitively assign performance to one cognitive trait, one way to validate that task structure is more likely to elicit performance based on the target cognitive trait is to assess the temporal and contextual repeatability of performance. In other words, individual performance is likely to represent an inherent trait when it is consistent across time and across similar or different tasks that theoretically test the same trait. Here, we assessed the temporal and contextual repeatability of performance on tasks intended to test the cognitive trait behavioral flexibility in great-tailed grackles (Quiscalus mexicanus). For temporal repeatability, we quantified the number of trials to form a color preference after each of multiple color reversals on a serial reversal learning task. For contextual repeatability, we then compared performance on the serial color reversal task to the latency to switch among solutions on each of two different multi-access boxes. We found that the number of trials to form a preference in reversal learning was repeatable across serial color reversals and the latency to switch a preference was repeatable across color reversal learning and the multi-access box contexts. This supports the idea that the reversal learning task structure elicits performance reflective of an inherent trait, and that reversal learning and solution switching on multi-access boxes similarly reflect the inherent trait of behavioral flexibility.

The role of climate change and niche shifts in divergent range dynamics of a sister-species pair.

Species ranges are set by limitations in factors including climate tolerances, habitat use, and dispersal abilities. Understanding the factors governing species range dynamics remains a challenge that is ever more important in our rapidly changing world. Species ranges can shift if environmental changes affect available habitat, or if the niche or habitat connectivity of a species changes. We tested how changes in habitat availability, niche, or habitat connectivity could contribute to divergent range dynamics in a sister-species pair. The great-tailed grackle (Quiscalus mexicanus) has expanded its range northward from Texas to Nebraska in the past 40 years, while its closest relative, the boattailed grackle (Quiscalus major), has remained tied to the coasts of the Atlantic Ocean and the Gulf of Mexico as well as the interior of Florida. We created species distribution and connectivity models trained on citizen science data from 1970-1979 and 2010-2019 to determine how the availability of habitat, the types of habitat occupied, and range-wide connectivity have changed for both species. We found that the two species occupy distinct habitats and that the great-tailed grackle has shifted to occupy a larger breadth of urban, arid environments farther from natural water sources. Meanwhile, the boattailed grackle has remained limited to warm, wet, coastal environments. We found no evidence that changes in habitat connectivity affected the ranges of either species. Overall, our results suggest that the great-tailed grackle has shifted its realized niche as part of its rapid range expansion, while the range dynamics of the boat-tailed grackle may be shaped more by climate change. The expansion in habitats occupied by the great-tailed grackle is consistent with observations that species with high behavioral flexibility can rapidly expand their geographic range by using human-altered habitat. This investigation identifies how opposite responses to anthropogenic change could drive divergent range dynamics, elucidating the factors that have and will continue to shape species ranges.

Great-tailed Grackles (Quiscalus mexicanus) as a tolerant host of avian malaria parasites.

Great-tailed Grackles (Quiscalus mexicanus) are a social, polygamous bird species whose populations have rapidly expanded their geographic range across North America over the past century. Before 1865, Great-tailed Grackles were only documented in Central America, Mexico, and southern Texas in the USA. Given the rapid northern expansion of this species, it is relevant to study its role in the dynamics of avian blood parasites. Here, 87 Great-tailed grackles in Arizona (a population in the new center of the range) were screened for haemosporidian parasites using microscopy and PCR targeting the parasite mitochondrial cytochrome b gene. Individuals were caught in the wild from January 2018 until February 2020. Haemosporidian parasite prevalence was 62.1% (54/87). A high Plasmodium prevalence was found (60.9%, 53/87), and one grackle was infected with Haemoproteus (Parahaemoproteus) sp. (lineage SIAMEX01). Twenty-one grackles were infected with P. cathemerium, sixteen with P. homopolare, four with P. relictum (strain GRW04), and eleven with three different genetic lineages of Plasmodium spp. that have not been characterized to species level (MOLATE01, PHPAT01, and ZEMAC01). Gametocytes were observed in birds infected with three different Plasmodium lineages, revealing that grackles are competent hosts for some parasite species. This study also suggests that grackles are highly susceptible and develop chronic infections consistent with parasite tolerance, making them competent to transmit some generalist haemosporidian lineages. It can be hypothesized that, as the Great-tailed Grackle expands its geographic range, it may affect local bird communities by increasing the transmission of local parasites but not introducing new species into the parasite species pool.

Using touchscreen equipped operant chambers to study animal cognition. Benefits, limitations, and advice.

Operant chambers are small enclosures used to test animal behavior and cognition. While traditionally reliant on simple technologies for presenting stimuli (e.g., lights and sounds) and recording responses made to basic manipulanda (e.g., levers and buttons), an increasing number of researchers are beginning to use Touchscreen-equipped Operant Chambers (TOCs). These TOCs have obvious advantages, namely by allowing researchers to present a near infinite number of visual stimuli as well as increased flexibility in the types of responses that can be made and recorded. We trained wild-caught adult and juvenile great-tailed grackles (Quiscalus mexicanus) to complete experiments using a TOC. We learned much from these efforts, and outline the advantages and disadvantages of our protocols. Our training data are summarized to quantify the variables that might influence participation and success, and we discuss important modifications to facilitate animal engagement and participation in various tasks. Finally, we provide a "training guide" for creating experiments using PsychoPy, a free and open-source software that was incredibly useful during these endeavors. This article, therefore, should serve as a resource to those interested in switching to or maintaining a TOC, or who similarly wish to use a TOC to test the cognitive abilities of non-model species or wild-caught individuals.

We can shift academic culture through publishing choices.

Researchers give papers for free (and often actually pay) to exploitative publishers who make millions off of our articles by locking them behind paywalls. This discriminates not only against the public (who are usually the ones that paid for the research in the first place), but also against the academics from institutions that cannot afford to pay for journal subscriptions and the 'scholarly poor'. I explain exploitative and ethical publishing practices, highlighting choices researchers can make right now to stop exploiting ourselves and discriminating against others.

Eurasian jays do not copy the choices of conspecifics, but they do show evidence of stimulus enhancement.

Corvids (birds in the crow family) are hypothesised to have a general cognitive tool-kit because they show a wide range of transferrable skills across social, physical and temporal tasks, despite differences in socioecology. However, it is unknown whether relatively asocial corvids differ from social corvids in their use of social information in the context of copying the choices of others, because only one such test has been conducted in a relatively asocial corvid. We investigated whether relatively asocial Eurasian jays (Garrulus glandarius) use social information (i.e., information made available by others). Previous studies have indicated that jays attend to social context in their caching and mate provisioning behaviour; however, it is unknown whether jays copy the choices of others. We tested the jays in two different tasks varying in difficulty, where social corvid species have demonstrated social information use in both tasks. Firstly, an object-dropping task was conducted requiring objects to be dropped down a tube to release a food reward from a collapsible platform, which corvids can learn through explicit training. Only one rook and one New Caledonian crow have learned the task using social information from a demonstrator. Secondly, we tested the birds on a simple colour discrimination task, which should be easy to solve, because it has been shown that corvids can make colour discriminations. Using the same colour discrimination task in a previous study, all common ravens and carrion crows copied the demonstrator. After observing a conspecific demonstrator, none of the jays solved the object-dropping task, though all jays were subsequently able to learn to solve the task in a non-social situation through explicit training, and jays chose the demonstrated colour at chance levels. Our results suggest that social and relatively asocial corvids differ in social information use, indicating that relatively asocial species may have secondarily lost this ability due to lack of selection pressure from an asocial environment.

Behavioral flexibility in an invasive bird is independent of other behaviors.

Behavioral flexibility is considered important for a species to adapt to environmental change. However, it is unclear how behavioral flexibility works: it relates to problem solving ability and speed in unpredictable ways, which leaves an open question of whether behavioral flexibility varies with differences in other behaviors. If present, such correlations would mask which behavior causes individuals to vary. I investigated whether behavioral flexibility (reversal learning) performances were linked with other behaviors in great-tailed grackles, an invasive bird. I found that behavioral flexibility did not significantly correlate with neophobia, exploration, risk aversion, persistence, or motor diversity. This suggests that great-tailed grackle performance in behavioral flexibility tasks reflects a distinct source of individual variation. Maintaining multiple distinct sources of individual variation, and particularly variation in behavioral flexibility, may be a mechanism for coping with the diversity of novel elements in their environments and facilitate this species' invasion success.

How far will a behaviourally flexible invasive bird go to innovate?

Behavioural flexibility is considered a key factor in the ability to adapt to changing environments. A traditional way of characterizing behavioural flexibility is to determine whether individuals invent solutions to novel problems, termed innovativeness. Great-tailed grackles are behaviourally flexible in that they can change their preferences when a task changes using existing behaviours; however, it is unknown how far they will go to invent solutions to novel problems. To begin to answer this question, I gave grackles two novel tests that a variety of other species can perform: stick tool use and string pulling. No grackle used a stick to access out-of-reach food, even after seeing a human demonstrate the solution. No grackle spontaneously pulled a vertically oriented string, but one did pull a horizontally oriented string twice. Additionally, a third novel test was previously conducted on these individuals and it was found that no grackle spontaneously dropped stones down a platform apparatus to release food, but six out of eight did become proficient after training. These results support the idea that behavioural flexibility is a multi-faceted trait because grackles are flexible, but not particularly innovative. This contradicts the idea that behavioural flexibility and innovativeness are interchangeable terms.

Behavioral flexibility and problem solving in an invasive bird.

Behavioral flexibility is considered an important trait for adapting to environmental change, but it is unclear what it is, how it works, and whether it is a problem solving ability. I investigated behavioral flexibility and problem solving experimentally in great-tailed grackles, an invasive bird species and thus a likely candidate for possessing behavioral flexibility. Grackles demonstrated behavioral flexibility in two contexts, the Aesop's Fable paradigm and a color association test. Contrary to predictions, behavioral flexibility did not correlate across contexts. Four out of 6 grackles exhibited efficient problem solving abilities, but problem solving efficiency did not appear to be directly linked with behavioral flexibility. Problem solving speed also did not significantly correlate with reversal learning scores, indicating that faster learners were not the most flexible. These results reveal how little we know about behavioral flexibility, and provide an immense opportunity for future research to explore how individuals and species can use behavior to react to changing environments.

Western scrub-jays do not appear to attend to functionality in Aesop's Fable experiments.

Western scrub-jays are known for their highly discriminatory and flexible behaviors in a caching (food storing) context. However, it is unknown whether their cognitive abilities are restricted to a caching context. To explore this question, we tested scrub-jays in a non-caching context using the Aesop's Fable paradigm, where a partially filled tube of water contains a floating food reward and objects must be inserted to displace the water and bring the food within reach. We tested four birds, but only two learned to drop stones proficiently. Of these, one bird participated in 4/5 experiments and one in 2/5 experiments. Both birds passed one experiment, but without attending to the functional differences of the objects, and failed the other experiments. Scrub-jays were not motivated to participate in these experiments, suggesting that either this paradigm was ecologically irrelevant or perhaps their flexibility is restricted to a caching context.

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.

Can endocranial volume be estimated accurately from external skull measurements in great-tailed grackles (Quiscalus mexicanus)?

There is an increasing need to validate and collect data approximating brain size on individuals in the field to understand what evolutionary factors drive brain size variation within and across species. We investigated whether we could accurately estimate endocranial volume (a proxy for brain size), as measured by computerized tomography (CT) scans, using external skull measurements and/or by filling skulls with beads and pouring them out into a graduated cylinder for male and female great-tailed grackles. We found that while females had higher correlations than males, estimations of endocranial volume from external skull measurements or beads did not tightly correlate with CT volumes. We found no accuracy in the ability of external skull measures to predict CT volumes because the prediction intervals for most data points overlapped extensively. We conclude that we are unable to detect individual differences in endocranial volume using external skull measurements. These results emphasize the importance of validating and explicitly quantifying the predictive accuracy of brain size proxies for each species and each sex.

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.

Validating methods for estimating endocranial volume in individual red deer (Cervus elaphus).

Comparing brain sizes is a key method in comparative cognition and evolution. Brain sizes are commonly validated by interspecific comparisons involving animals of varying size, which does not provide a realistic index of their accuracy for intraspecific comparisons. Intraspecific validation of methods for measuring brain size should include animals of the same age and sex to ensure that individual differences can be detected in animals of similar size. In this study we compare three methods of measuring the endocranial volume of 33 red deer skulls to investigate the accuracy of each method. Methods for estimating endocranial volume included scanning each skull using computerised tomography (CT) and quantifying the volume with OsiriX software, filling the cranium with glass beads and measuring the bead volume, and linear measurements (length, width, and height) of the cranium using callipers. CT scan volumes were highly correlated with results from the bead method, but only moderately correlated with the linear method. This study illustrates the importance of validating intraspecies measurement methods, which allows for the accurate interpretation of results.

Specializations of birds that attend army ant raids: an ecological approach to cognitive and behavioral studies.

Tropical birds forage at army ant raids on several continents. Obligate foraging at army ant raids evolved several times in the Neotropical true antbird family (Thamnophilidae), and recent evidence suggests a diversity of bird species from other families specialize to varying degrees on army ant exploitation. Army ant raids offer access to high prey densities, but the ant colonies are mobile and widely spaced. Successful army ant exploitation requires solving a complex foraging problem because army ant raids are unpredictable in space and time. Birds can counteract the challenges posed by the ants by using strategies that raise their chances of detecting army ant raids, and birds can use additional strategies to track army ant colonies they have located. Some features of army ant biology, such as their conspicuous swarms and columns, above-ground activity, and regular cycles of behavior, provide opportunities for birds to increase their effectiveness at exploiting raids. Changes in sensory, cognitive and behavioral systems may all contribute to specialized army ant exploitation in a bird population. The combination of specializations that are employed may vary independently among bird species and populations. The degree of army ant exploitation by birds varies geographically with latitude and elevation, and with historical patterns such as centers of distribution of obligate thamnophilid antbirds. We predict the set of specializations a given bird population exhibits will depend on local ecology, as well as phylogenetic history. Comparative approaches that focus on these patterns may indicate ecological and evolutionary factors that have shaped the costs and benefits of this foraging strategy. The development of army ant exploitation in individual birds is poorly understood, and individual expression of these specializations may depend on a combination of genetic adaptation with cognitive plasticity, possibly including social and experiential learning. Future studies that measure developmental changes and quantify individual differences in army ant exploitation are needed to establish the mechanisms underlying this behavior.