Alan B. Bond

Pinyon jays use transitive inference to predict social dominance

Living in large, stable social groups is often considered to favour the evolution of enhanced cognitive abilities, such as recognizing group members, tracking their social status and inferring relationships among them. An individuals place in the social order can be learned through direct interactions with others, but conflicts can be time-consuming and even injurious. Because the number of possible pairwise interactions increases rapidly with group size, members of large social groups will benefit if they can make judgments about relationships on the basis of indirect evidence. Transitive reasoning should therefore be particularly important for social individuals, allowing assessment of relationships from observations of interactions among others. Although a variety of studies have suggested that transitive inference may be used in social settings, the phenomenon has not been demonstrated under controlled conditions in animals. Here we show that highly social pinyon jays (Gymnorhinus cyanocephalus) draw sophisticated inferences about their own dominance status relative to that of strangers that they have observed interacting with known individuals. These results directly demonstrate that animals use transitive inference in social settings and imply that such cognitive capabilities are widespread among social species.

Social complexity and transitive inference in corvids

The social complexity hypothesis asserts that animals living in large social groups should display enhanced cognitive abilities along predictable dimensions. To test this concept, we compared highly social pinyon jays, Gymnorhinus cyanocephalus, with relatively nonsocial western scrub-jays, Aphelocoma californica, on two complex cognitive tasks relevant to the ability to track and assess social relationships. Pinyon jays learned to track multiple dyadic relationships more rapidly and more accurately than scrub-jays and appeared to display a more robust and accurate mechanism of transitive inference. These results provide a clear demonstration of the association between social complexity and cognition in animals.

Visual predators select for crypticity and polymorphism in virtual prey

Cryptically coloured animals commonly occur in several distinct pattern variants. Such phenotypic diversity may be promoted by frequency-dependent predation, in which more abundant variants are attacked disproportionately often, but the hypothesis has never been explicitly tested. Here we report the first controlled experiment on the effects of visual predators on prey crypticity and phenotypic variance, in which blue jays (Cyanocitta cristata) searched for digital moths on computer monitors. Moth phenotypes evolved via a genetic algorithm in which individuals detected by the jays were much less likely to reproduce. Jays often failed to detect atypical cryptic moths, confirming frequency-dependent selection and suggesting the use of searching images, which enhance the detection of common prey. Over successive generations, the moths evolved to become significantly harder to detect, and they showed significantly greater phenotypic variance than non-selected or frequency-independent selected controls.

Optimal Foraging in a Uniform Habitat: The Search Mechanism of the Green Lacewing

Abstract The effects of food deprivation and prey contact on the components of searching behaviour in larval green lacewings ( Chrysopa carnea Stephens) were examined to test the applicability of optimal foraging theory to predation in a uniform habitat. Variation in foraging intensity was primarily the result of changes in the meander. Modulation of the response to prey contact with increasing deprivation involved changes in the velocity and the response persistence and suggested the occurrence of adaptation to inferred differences in the spatial distribution of the prey. The ratio of giving-up times at different levels of deprivation was in accordance with the predictions of optimal foraging theory, even though the conditions precluded a discrete decision process.

Apostatic selection by blue jays produces balanced polymorphism in virtual prey

Apostatic selection, in which predators overlook rare prey types while consuming an excess of abundant ones, has been assumed to contribute to the maintenance of prey polymorphisms. Such an effect requires predators to respond to changes in the relative abundance of prey, switching to alternatives when a focal prey type becomes less common,. Apostatic selection has often been investigated using fixed relative proportions of prey,, but its effects on predator–prey dynamics have been difficult to demonstrate. Here we report results from a new technique that incorporates computer-generated displays, into an established experimental system, that of blue jays (Cyanocitta cristata) hunting for cryptic Catocala moths. Digital prey images from a virtual population are presented to predators. The relative numbers that escape detection determine the subsequent abundance of each prey type. If apostatic selection does promote stability, the system should converge on an eqlibrium in which each prey type appears at a characteristic abundance. Our results show that the detection of cryptic prey does involve apostatic selection, and that such selection can function to maintain prey polymorphism.

Spatial memory and the performance of rats and pigeons in the radial-arm maze

The resource-distribution hypothesis states that the ability of an animal to remember the spatial location of past events is related to the typical distribution of food resources for the species. It appears to predict that Norway rats would perform better than domestic pigeons in tasks requiring spatial event memory. Pigeons, tested in an eight-arm radial maze, exhibited no more than half of the memory capacity observed in rats in the same apparatus and may not have used spatial memory at all. The results were interpreted as supporting the hypothesis.

Serial Reversal Learning and the Evolution of Behavioral Flexibility in Three Species of North American Corvids (Gymnorhinus cyanocephalus, Nucifraga columbiana, Aphelocoma californica)

In serial reversal learning, subjects learn to respond differentially to 2 stimuli. When the task is fully acquired, reward contingencies are reversed, requiring the subject to relearn the altered associations. This alternation of acquisition and reversal can be repeated many times, and the ability of a species to adapt to this regimen has been considered as an indication of behavioral flexibility. Serial reversal learning of 2-choice discriminations was contrasted in 3 related species of North American corvids: pinyon jays (Gymnorhinus cyanocephalus), which are highly social; Clarks nutcrackers (Nucifraga columbiana), which are relatively solitary but specialized for spatial memory; and western scrub jays (Aphelocoma californica), which are ecological generalists. Pinyon jays displayed significantly lower error rates than did nutcrackers or scrub jays after reversal of reward contingencies for both spatial and color stimuli. The effect was most apparent in the 1st session following each reversal and did not reflect species differences in the rate of initial discrimination learning. All 3 species improved their performance over successive reversals and showed significant transfer between color and spatial tasks, suggesting a generalized learning strategy. The results are consistent with an evolutionary association between behavioral flexibility and social complexity.

A Comparative Analysis of Social Play in Birds

Summary Although social play is broadly distributed among mammals, it is infrequently encountered in other vertebrate taxa. It is, however, displayed in a fully realized and complex form in several groups of birds. Unambiguous accounts of social play have been recorded from thirteen species of parrots, seven species of corvids, and several hornbills and Eurasian babblers. We conducted an analysis of the avian play literature, testing for differences between avian taxa, as well as for correlations between play complexity, brain size, and age of first reproduction. Corvids were far more likely to show social object play than parrots. Corvids, parrots, and hornbills had larger relative brain sizes than would be predicted from a class-level allometric regression, but brain size was not associated with the complexity of social play among genera within taxa. Play complexity within parrots and corvids was, however, significantly associated with the age of first re production. The likelihood of complex social play appears to increase when delayed reproduction is accompanied by persisting relationships between adults and post-fledging juveniles. The adaptive significance of social play in birds thus offers intriguing parallels to similar analyses in mammals.

The foraging behaviour of lacewing larvae on vertical rods

Abstract The foraging behaviour of lacewing larvae (Chrysopa carnea Stephens, Chrysopidae, Neuroptera) on vertical lucite rods was observed under a variety of experimental conditions to investigate the decision processes responsible for the distribution of foraging effort. Food deprivation increased the duration of searching on all parts of the rod, whereas contact with prey at the rod tip induced only a local enhancement of searching activity. Searching at the rod tip did not decline with repeated trials on the same rod, but the duration of searching on the rest of the rod was reduced, evidently reflecting recognition and avoidance of previously-searched substrates. There were indications that the larvae compensated for changes in the surface area, radius of curvature, and length of the rod. Searching patterns at the tip and on the rest of the rod proved to be independently modifiable, suggesting a two-tiered hierarchy of foraging decisions. The total duration of the search depended primarily on the pattern of vertical movement and the number of reascents, rather than the searching effort per unit area. Evidence of a negative geotaxis was found only during the initial ascent of the rod, and no sequential pattern in the lengths of vertical excursions was observed. The decision process appeared to involve a combination of variable control factors interacting with a generally constant, random probability of change or reversal of direction. The resulting investment of foraging time corresponded surprisingly well with expectations derived from optimal foraging theory, thereby testifying to the power of approximate, semi-random decision processes.

Searching image in blue jays: Facilitation and interference in sequential priming

Repeated exposure to a single target type (sequential priming) during visual search for multiple cryptic targets commonly improves performance on subsequent presentations of that target. It appears to be an attentional phenomenon, a component of the searching image effect. It has been argued, however, that if searching image is an attentional process, sequential priming should also interfere with performance on subsequent nonprimed targets, and such interference has never been unequivocally demonstrated. In blue jays (Cyanocitta cristata) searching in an operant apparatus for targets derived from images of cryptic moths, detection performance was strongly facilitated in the course of a sequential prime but was relatively unaffected by sequences of mixed target types. Detection accuracy in subsequent probe trials was enhanced by priming with targets of the same type, whereas accuracy on cryptic probes following priming with a more conspicuous target was significantly degraded. The results support an attentional interpretation of searching image.