Ecological factors shape quantitative decision-making in coyotes.

Much research has focused on the development and evolution of cognition in the realm of numerical knowledge in human and nonhuman animals but often fails to take into account ecological realities that, over time, may influence and constrain cognitive abilities in real-life decision-making. Cognitive abilities such as enumerating and timing are central to many psychological and ecological models of behavior, yet our knowledge of how these are affected by environmental fluctuations remains incomplete. Our research bridges the gap between basic cognitive research and ecological decision-making. We used coyotes (Canis latrans) as a model animal system to study decision-making about smaller, more proximal food rewards and larger, more distant food rewards; we tested animals across their four reproductive cycle phases to examine effects of ecological factors such as breeding status and environmental risk on quantitative performance. Results show that coyotes, similar to other species, spatially discount food rewards while foraging. The degree to which coyotes were sensitive to the risk of obtaining the larger food reward, however, depended on the season in which they completed the foraging task, the presence of unfamiliar humans (i.e., risk), and the presence of conspecifics. Importantly, our results support that seasonal variations drive many differences in nonhuman animal behavior and cognition (e.g., hibernation, breeding, food resource availability). Further, it may be useful in the future to extend this work to humans because seasons may influence human cognition as well, and this remains unexplored in the realms of enumeration, timing, and spatial thinking.

Persistence and conspecific observations improve problem-solving abilities of coyotes.

Social learning has important ecological and evolutionary consequences but the role of certain factors, such as social rank, neophobia (i.e., avoidance of novel stimuli), persistence, and task-reward association, remain less understood. We examined the role of these factors in social learning by captive coyotes (Canis latrans) via three studies. Study 1 involved individual animals and eliminated object neophobia by familiarizing the subjects to the testing apparatus prior to testing. Studies 2 and 3 used mated pairs to assess social rank, and included object neophobia, but differed in that study 3 decoupled the food reward from the testing apparatus (i.e., altered task-reward association). For all three studies, we compared performance between coyotes that received a demonstration from a conspecific to control animals with no demonstration prior to testing. Coyotes displayed social learning during study 1; coyotes with a demonstrator were faster and more successful at solving the puzzle box but did not necessarily use the same modality as that observed to be successful. In study 2, there was no difference in success between treatment groups but this is likely because only one coyote within each pair was successful so successful coyote results were masked by their unsuccessful mate. In study 3, there was no difference in success between treatment groups; only two coyotes, both dominant, hand-reared males with demonstrators were able to perform the task. However, coyotes with a demonstrator were less neophobic, measured as latency to approach the object, and more persistent, measured as time spent working on the apparatus. Social rank was the best predictor of neophobia and persistence and was also retained in the best model for time to eat inside the apparatus, a post-trial measurement of object neophobia. These results suggest coyotes are capable of social learning for novel tasks but social rank, neophobia, and persistence influence their social-learning capabilities. This study contributes to understanding the mechanisms underlying how animals gain information about their environment.

The intrepid urban coyote: a comparison of bold and exploratory behavior in coyotes from urban and rural environments.

Coyotes (Canis latrans) are highly adaptable, medium-sized carnivores that now inhabit nearly every large city in the United States and Canada. To help understand how coyotes have adapted to living in urban environments, we compared two ecologically and evolutionarily important behavioral traits (i.e., bold-shy and exploration-avoidance behavior) in two contrasting environments (i.e., rural and urban). Boldness is an individual's reaction to a risky situation and exploration is an individual's willingness to explore novel situations. Our results from both tests indicate that urban coyotes are bolder and more exploratory than rural coyotes and that within both populations there are individuals that vary across both spectrums. Bolder behavior in urban coyotes emerged over several decades and we speculate on possible processes (e.g., learning and selection) and site differences that could be playing a role in this behavioral adaptation. We hypothesize that an important factor is how people treat coyotes; in the rural area coyotes were regularly persecuted whereas in the urban area coyotes were rarely persecuted and sometimes positively rewarded to be in close proximity of people. Negative consequences of this behavioral adaptation are coyotes that become bold enough to occasionally prey on pets or attack humans.

Coyotes (Canis latrans) and the matching law.

Environmental change is accelerating due to anthropogenic influence. Species that have greater behavioral flexibility may be better adapted to exploit new or constantly changing habitats. There are few mammals and even fewer carnivores that better illustrate widespread adaptability and behavioral flexibility in the wake of human disturbance than coyotes (Canis latrans). Yet how such predators successfully track resources, enabling them to survive and extend their range in stochastic environments remains unknown. We tested eight wild-born, captive coyotes individually on an operant two-choice test using concurrent variable interval (VI) schedules. We held the overall rate of reinforcement constant but manipulated the ratio of reinforcement available from the two choices. We analyzed sensitivity of coyotes' tracking of resource change by fitting the generalized matching equation to the data. Results showed all coyotes efficiently tracked changes in reinforcement ratios within the first few sessions of each new condition and matched their relative rate of foraging time to relative rate of resources. We suggest the matching paradigm provides a methodology to explore coyote foraging strategies, and a potential framework to compare behavioral flexibility across species, by measuring the ability to track resource change under variable resource conditions.

Characterizing noise in nonhuman vocalizations: Acoustic analysis and human perception of barks by coyotes and dogs.

Measuring noise as a component of mammalian vocalizations is of interest because of its potential relevance to the communicative function. However, methods for characterizing and quantifying noise are less well established than methods applicable to harmonically structured aspects of signals. Using barks of coyotes and domestic dogs, we compared six acoustic measures and studied how they are related to human perception of noisiness. Measures of harmonic-to-noise-ratio (HNR), percent voicing, and shimmer were found to be the best predictors of perceptual rating by human listeners. Both acoustics and perception indicated that noisiness was similar across coyote and dog barks, but within each species there was significant variation among the individual vocalizers. The advantages and disadvantages of the various measures are discussed.