Corina Jill Logan

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.

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.

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.

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.

Beyond Brain Size

Despite prolonged interest in comparing brain size and behavioral proxies of ‘intelligence’ across taxa, the adaptive and cognitive significance of brain size variation remains elusive. Central to this problem is the continued focus on hominid cognition as a benchmark, and the assumption that behavioral complexity has a simple relationship with brain size. Although comparative studies of brain size have been criticized for not reflecting how evolution actually operates, and for producing spurious, inconsistent results, the causes of these limitations have received little discussion. We show how these issues arise from implicit assumptions about what brain size measures and how it correlates with behavioral and cognitive traits. We explore how inconsistencies can arise through heterogeneity in evolutionary trajectories and selection pressures on neuroanatomy or neurophysiology across taxa. We examine how interference from ecological and life history variables complicates interpretations of brain-behavior correlations, and point out how this problem is exacerbated by the limitations of brain and cognitive measures. These considerations, and the diversity of brain morphologies and behavioral capacities, suggest that comparative brain-behavior research can make greater progress by focusing on specific neuroanatomical and behavioral traits within relevant ecological and evolutionary contexts. We suggest that a synergistic combination of the ‘bottom up’ approach of classical neuroethology and the ‘top down’ approach of comparative biology/psychology within closely related but behaviorally diverse clades can limit the effects of heterogeneity, interference, and noise. We argue this shift away from broad-scale analyses of superficial phenotypes will provide deeper, more robust insights into brain evolution.

Innovation does not indicate behavioral flexibility in great-tailed grackles

Many cross-species studies attest that innovation frequency (novel food types eaten and foraging techniques used) is a measure of behavioral flexibility and show that it positively correlates with relative brain size (corrected for body size). However, mixed results from the three studies that directly test the relationship between innovation frequency and behavioral flexibility and behavioral flexibility and brain size question both assumptions. I investigated behavioral flexibility in non-innovative great-tailed grackles that have an average sized brain, and compared their test performance with innovative, large-brained New Caledonian crows. Contrary to the prediction, grackles perform similarly to crows in experiments using clear tubes partially filled with water and containing a floating food reward, where objects must be dropped into the tube to raise the water level, bringing the food within reach. Similarly to crows, 4 out of 6 grackles preferred to drop the more functional heavy (rather than light) objects, and 2 changed their preference in a follow up experiment where the heavy objects were no longer functional, thus exhibiting behavioral flexibility. These results challenge the assumption that innovation frequency indicates behavioral flexibility since a non-innovative bird demonstrated behavioral flexibility at a level similar to that in innovative crows, and they challenge the assumption that only large brains are capable of behavioral flexibility because a bird with an average brain size solved problems similarly to large-brained crows.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 abilities experimentally in great-tailed grackles, an invasive species and thus a likely candidate for possessing behavioral flexibility. I found that grackles are behaviorally flexible and good problem solvers, they vary in behavioral flexibility across contexts, flexibility did not correlate with problem solving ability, and those that are more flexible did not necessarily use more learning strategies. It appears that behavioral flexibility can be an independent trait that varies across contexts. Maintaining such a high level of variation could be a mechanism underlying successful species invasions. These results highlight the need to investigate how individuals use behavior to react to changing environments.

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.

Convergent minds: the evolution of cognitive complexity in nature

Stephen Jay Gould [[1][1],[2][2]] argued that replaying the ‘tape of life’ would result in radically different evolutionary outcomes. Goulds ‘radical contingency thesis’ focused primarily on animal morphology, and in particular on the specific parameters of animal body plans, which, he

Endocranial volume is heritable and is associated with longevity and fitness in a wild mammal

Research on relative brain size in mammals suggests that increases in brain size may generate benefits to survival and costs to fecundity: comparative studies of mammals have shown that interspecific differences in relative brain size are positively correlated with longevity and negatively with fecundity. However, as yet, no studies of mammals have investigated whether similar relationships exist within species, nor whether individual differences in brain size within a wild population are heritable. Here we show that, in a wild population of red deer (Cervus elaphus), relative endocranial volume was heritable (h2 = 63%; 95% credible intervals (CI) = 50–76%). In females, it was positively correlated with longevity and lifetime reproductive success, though there was no evidence that it was associated with fecundity. In males, endocranial volume was not related to longevity, lifetime breeding success or fecundity.

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.