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Scale-dependent effects of density and habitat on foal survival
Identifying the most appropriate scale to study factors influencing life history is important to evolutionary ecology and wildlife management. For example, the scale at which density is assessed and explains variation in survival can affect how biologists observe and interpret population dynamics, which can influence plans for managing populations. Feral horses (Equus ferus caballus) contrast with most ungulates by exhibiting a mating system characterized by female-defense polygyny with persistent, non-territorial breeding groups (bands) and female-biased initial (natal) and subsequent (breeding) dispersal. We predicted that for horses, offspring movements coupled with female-biased breeding dispersal would increase the scale at which density best related to juvenile survival compared to species with greater female philopatry. From 2008 to 2013, we censused the population of feral horses on Sable Island, Canada. We annually computed individual-specific local densities for 442 foals (horses/km2 in radii of 2,000 m, 4,000 m, and 8,000 m fixed to a band’s centroid of movements) and whole-island (total) population density, group (band) size, and local access to surface freshwater, which affected movement patterns and selection of vegetation by females. The population of feral horses increased from 380 in 2008 to 559 in 2013. Overwinter survival of foals averaged 82.8%. Island-wide density was the most important predictor of foal mortality and was negatively associated with survival, with a lesser negative effect from local density. Increased access to surface freshwater (ponds) was an important predictor of foal survival but only at certain scales. Our study emphasizes the relevance of a multi-scale approach when analyzing the response of fitness components to changes in habitat and population processes, which may be influenced by the particular social organization of the species
Island tameness and the repeatability of flight initiation distance in a large herbivore
This is the author accepted manuscript. The final version is available from NRC Research Press via the DOI in this record.Antipredator behaviours can be lost relatively quickly in populations that are relieved of predation, as is known for several species inhabiting islands. Flight initiation distance (FID) is often studied in the context of island tameness; however, little is known about the factors that influence and maintain FID variation in predation-free populations. Here, we studied FID in foals of an isolated predator-free population of feral horses (Equus caballus L., 1758) on Sable Island, Canada, to determine if FID could be used for research on consistent individual differences in risk aversion and island tameness. In addition to testing for temporal, spatial, and sex effects on FID, we compared repeatability estimates at two temporal scales (within and among days). Similar FID for measurements obtained on the same day and for males and females indicated an absence of short-term desensitization and sex effects. In contrast, FID decreased for measurements made on subsequent days and from east to west, which could reflect habituation to human presence and (or) other temporal and spatial processes. Repeatability was high (0.42 ± 0.06), but tended to decrease with increasing time intervals. This study highlights the potential of FID for individual-based research on the ecology and evolutionary dynamics of risk aversion in predation-free populations.Funding was provided by the Natural Sciences and Engineering Research Council of Canada (NSERC Discovery Grant No. 371535-2009 to P.D.M.), the Canada Foundation for Innovation (Leaders Opportunity Grant No. 25046 to P.D.M.), and a Royal Society International Exchange grant (J.P. and P.D.M.). D.A. was supported by an NSERC Ph.D. scholarship. D.C. received support from the University of Exeter M.Sc. program in Evolutionary and Behavioural Ecology. J.P. was supported by a Leverhulme Trust Early Career Research Fellowship
The ontogeny of antipredator behavior: age differences in California ground squirrels (Otospermophilus beecheyi) at multiple stages of rattlesnake encounters
Newborn offspring of animals often exhibit fully functional innate antipredator behaviors, but they may also require learning or further development to acquire appropriate responses. Experience allows offspring to modify responses to specific threats and also leaves them vulnerable during the learning period. However, antipredator behaviors used at one stage of a predator encounter may compensate for deficiencies at another stage, a phenomenon that may reduce the overall risk of young that are vulnerable at one or more stages. Few studies have examined age differences in the effectiveness of antipredator behaviors across multiple stages of a predator encounter. In this study, we examined age differences in the antipredator behaviors of California ground squirrels (Otospermophilus beecheyi) during the detection, interaction, and attack stages of Pacific rattlesnake (Crotalus oreganus) encounters. Using free-ranging squirrels, we examined the ability to detect free-ranging rattlesnakes, snake-directed behaviors after discovery of a snake, and responses to simulated rattlesnake strikes. We found that age was the most important factor in snake detection, with adults being more likely to detect snakes than pups. We also found that adults performed more tail flagging (a predator-deterrent signal) toward snakes and were more likely to investigate a snake’s refuge when interacting with a hidden snake. In field experiments simulating snake strikes, adults exhibited faster reaction times than pups. Our results show that snake detection improves with age and that pups probably avoid rattlesnakes and minimize time spent in close proximity to them to compensate for their reduced reaction times to strikes