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ABSTRACT Cities impose unique selection pressures on wildlife and generate clines in phenotypic traits along urban–rural gradients. Roads are a widespread feature of human‐dominated landscapes and are known to cause direct wildlife mortality; however, whether they act as a selective force influencing phenotypic trait variation along urban–rural gradients remains unclear. This study tested the hypothesis that roads influence natural selection of coat color in the eastern gray squirrel (Sciurus carolinensis), a species with two distinct coat colors: a gray morph that is common in all areas and a melanic morph more prevalent in urban areas than in rural ones. Vehicular collisions are a significant cause of mortality in eastern gray squirrels, with the melanic morph more visually conspicuous on roads and more easily detected and avoided by drivers than the gray morph. Standardized road cruise surveys along an urbanization gradient in Syracuse, New York, USA, revealed that the prevalence of melanism among living squirrels in Syracuse was negatively related to distance from the city center, whereas there was no urban–rural cline in melanism among road‐killed individuals, with the melanic morph underrepresented among road‐killed squirrels by up to 30% along the urbanization gradient. An examination of the prevalence of each color morph on and off road surfaces in a range‐wide compilation of > 100,000 photographs ofS. carolinensisalso indicated that the melanic morph was underrepresented among road‐killed squirrels imaged. Our study highlights vehicular collisions as an important source of natural selection on phenotypic traits, suggesting a potential role in shaping patterns of urban evolution and contributing to the maintenance of urban–rural clines. 

Abstract Phenotypic differences between urban and rural populations are well‐documented, but the evolutionary processes driving trait variation along urbanization gradients are often unclear. We combined spatial data on abundance, trait variation, and measurements of fitness to understand cline structure and test for natural selection on heritable coat color morphs (melanic, gray) of eastern gray squirrels (Sciurus carolinensis) along an urbanization gradient. Population surveys using remote cameras and visual counts at 76 sites along the urbanization gradient revealed a significant cline in melanism, decreasing from 48% in the city center to <5% in rural woodlands. Among 76 squirrels translocated to test for phenotypic selection, survival was lower for the melanic than gray morph in rural woodlands, whereas there was no difference in survival between color morphs in the city. These results suggest the urban–rural cline in melanism is explained by natural selection favoring the gray morph in rural woodlands combined with relaxed selection in the city. Our study illustrates how trait variation between urban and rural populations can emerge from selection primarily in rural populations rather than adaptation to novel features of the urban environment. 

Abstract Shrub encroachment is transforming arid and semiarid grasslands worldwide. Such transitions should influence predator–prey interactions because vegetation cover often affects risk perception by prey and contributes to their landscape of fear. We examined how the landscape of fear of two desert lagomorphs (black‐tailed jackrabbit,Lepus californicus; desert cottontail,Sylvilagus audubonii) changes across grassland‐to‐shrubland gradients at Jornada Basin Long Term Ecological Research site in the Chihuahuan Desert of southern New Mexico. We test whether shrub encroachment shapes risk differently for these two lagomorphs because of differences in body size and predator escape tactics. We also examine whether an ecosystem engineer of grasslands (banner‐tailed kangaroo rat,Dipodomys spectabilis) mediates risk perception through the creation of escape refuge and whether trade‐offs exist between shrub encroachment and the local reduction of banner‐tailed kangaroo rats caused by shrub expansion. We measured perceived predation risk with flight initiation distances (FIDs) and then used structural equation modeling to tease apart the hypothesized direct and indirect pathways for how shrub encroachment could affect perceived risk. A total negative effect of shrub cover on FID was supported for jackrabbits and cottontails, suggesting both species perceive shrubbier habitat as safer. Increases in fine‐scale concealment also reduced risk for cottontails, but not jackrabbits, likely because cottontails rely on crypsis to avoid predator detection whereas jackrabbits rely on speed and agility to outrun predators. Perceived risk was reduced when individuals were near kangaroo rat mounds only for cottontails because the smaller species can use banner‐tailed kangaroo rat mounds as refuge. Shrub encroachment greatly reduced the availability of mounds. Thus, a trade‐off exists for cottontails in which shrub encroachment directly reduced perceived risk, but indirectly increased perceived risk through the local extirpation of an ecosystem engineer. Our work illustrates how the expansion of shrub encroachment can create a dynamic landscape of fear for populations of prey species involving direct and indirect pathways contingent on prey body size, escape tactics, and activities of an ecosystem engineer.