Effects of sun compass error on spatial search by Clark's nutcrackers.

Animals employ compasses during navigation, but little attention has been paid to how accuracy is maintained in the face of compass error, which is inevitable in biological systems. The use of multiple landmarks may minimize the effect of compass error. We allowed Clark's nutcrackers to cache seeds in an outdoor aviary with either one or four landmarks present, and subsequently subjected them to small clock-shifts mimicking the effects of compass error. As predicted, the results showed a significant decrease in search accuracy following the clock-shift when one landmark was present but not when four landmarks were present. These results support that nutcrackers encode information from the sun as well as terrestrial landmarks, and these spatial cues are used in a flexible manner. Overall, our results are important as they support the hypothesis that multiple landmarks may be used during situations where the sun compass has even a small amount of error.

Clark's nutcracker spatial memory: the importance of large, structural cues.

Clark's nutcrackers, Nucifraga columbiana, cache and recover stored seeds in high alpine areas including areas where snowfall, wind, and rockslides may frequently obscure or alter cues near the cache site. Previous work in the laboratory has established that Clark's nutcrackers use spatial memory to relocate cached food. Following from aspects of this work, we performed experiments to test the importance of large, structural cues for Clark's nutcracker spatial memory. Birds were no more accurate in recovering caches when more objects were on the floor of a large experimental room nor when this room was subdivided with a set of panels. However, nutcrackers were consistently less accurate in this large room than in a small experimental room. Clark's nutcrackers probably use structural features of experimental rooms as important landmarks during recovery of cached food. This use of large, extremely stable cues may reflect the imperfect reliability of smaller, closer cues in the natural habitat of Clark's nutcrackers. This article is part of a Special Issue entitled: CO3 2013.

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; Clark's 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.

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 individual's 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.

Discrimination among pinyon pine trees by Clark's Nutcrackers: effects of cone crop size and cone characters.

The influences of Colorado pinyon pine (Pinus edulis) cone crop size, cone and seed weight, cone length, number of seeds per cone, number of viable seeds, and percent viable seeds on the foraging behavior of avian seed dispersal agents were examined in field and laboratory settings. In the field, there was a significant positive relationship between cone number per tree and both the absolute number of cones and the percentage of the cone crop from which seeds were harvested. Cone weight and the number of viable seeds were also significantly related to seed harvest intensity. Laboratory experiments examined the relationship between crop size and cone characters on seed harvest by 18 Clark's Nutcrackers (Nucifraga columbiana). Nutcrackers were offered a choice of two tree types: one with 20 cones attached, and another with 10 cones attached. Significantly more birds chose to remove seeds first from the tree with 20 cones than the tree with 10 cones. In timed trials, they also harvested seeds from significantly more cones on the tree with the higher cone density. In the laboratory, cones chosen for seed removal by the nutcrackers had significantly more viable seeds, more seeds, and were longer compared to cones that were not chosen. Such discriminatory foraging behavior may increase avian foraging efficiency and result in differential reproductive success of pinyon pines. This behavior may therefore influence the evolution of pinyon pine reproductive traits.