The amount of risk animals perceive in a given circumstance (i.e. their degree of 'fear') is a difficult motivational state to study. While many studies have used flight initiation distance as a proxy for fearfulness and examined the factors influencing the decision to flee, there is no general understanding of the relative importance of these factors. By identifying factors with large effect sizes, we can determine whether anti-predator strategies reduce fear, and we gain a unique perspective on the coevolution of predator and anti-predator behaviour. Based on an extensive review and formal meta-analysis, we found that predator traits that were associated with greater risk (speed, size, directness of approach), increased prey distance to refuge and experience with predators consistently amplified the perception of risk (in terms of flight initiation distance). While fish tolerated closer approach when in larger schools, other taxa had greater flight initiation distances when in larger groups. The presence of armoured and cryptic morphologies decreased perception of risk, but body temperature in lizards had no robust effect on flight initiation distance. We find that selection generally acts on prey to be sensitive to predator behaviour, as well as on prey to modify their behaviour and morphology.

Weaponry is ubiquitous in male ungulates and is driven by intrasexual selection, but the mystery surrounding its sporadic presence in females remains unsolved. Female horns are often smaller and shaped differently to male horns, suggesting a different function; indeed, hypotheses explaining the presence of female horns include competition for food, male mollification and defence against predators. Here we use comparative phylogenetic analyses to show that females are significantly more likely to bear horns in bovids that are conspicuous due to large body size and living in open habitats than inconspicuous species living in closed habitats or that are small. An inability to rely on crypsis or take refuge in deep vegetation has apparently driven the evolution of horns for defence against predators in female bovids, a finding supported by many field observations. Typically, exceptions are small species where females are territorial (e.g. duikers) and use horns in intrasexual contests. Furthermore, we suggest that conspicuousness and territoriality hypotheses may explain other instances of female cranial weaponry (i.e. antlers and ossicones) in other horned ruminants. Our phylogenetic reconstruction indicates that the primary function of horns in females is linked to antipredator defence in most clades, but occasionally to intrasexual competition in others.

Understanding how biodiversity responds to urbanization is challenging, due in part to the single‐city focus of most urban ecological research. Here, we delineate continent‐scale patterns in urban species assemblages by leveraging data from a multi‐city camera trap survey and quantify how differences in greenspace availability and average housing density among 10 North American cities relate to the distribution of eight widespread North American mammals. To do so, we deployed camera traps at 569 sites across these ten cities between 18 June and 14 August. Most data came from 2017, though some cities contributed 2016 or 2018 data if it was available. We found that the magnitude and direction of most species' responses to urbanization within a city were associated with landscape‐scale differences among cities. For example, eastern gray squirrel (Sciurus carolinensis), fox squirrel (Sciurus niger), and red fox (Vulpes vulpes) responses to urbanization changed from negative to positive once the proportion of green space within a city was >~20%. Likewise, raccoon (Procyon lotor) and Virginia opossum (Didelphis virginiana) responses to urbanization changed from positive to negative once the average housing density of a city exceeded about 700 housing units/km2. We also found that local species richness within cities consistently declined with urbanization in only the more densely developed cities (>~700 housing units/km2). Given our results, it may therefore be possible to design cities to better support biodiversity and reduce the negative influence of urbanization on wildlife by, for example, increasing the amount of green space within a city. Additionally, it may be most important for densely populated cities to find innovative solutions to bolster wildlife resilience because they were the most likely to observe diversity losses of common urban species.

When a previously common predator disappears owing to local extinction, the strong source of natural selection on prey to visually recognize that predator becomes relaxed. At present, we do not know the extent to which recognition of a specific predator is generalized to similar looking predators or how a specific predator-recognition cue, such as coat pattern, degrades under prolonged relaxed selection. Using predator models, we show that deer exhibit a more rapid and stronger antipredator response to their current predator, the puma, than to a leopard displaying primitive rosettes similar to a locally extinct predator, an early jaguar. Presentation of a novel tiger with a striped coat engendered an intermediate speed of predator recognition and strength of antipredator behaviour. Responses to the leopard model slightly exceeded responses to a non-threatening deer model, suggesting that thousands of years of relaxed selection have led to the loss of recognition of the spotted coat as a jaguar-recognition cue, and that the spotted coat has regained its ability to camouflage the felid form. Our results shed light on the evolutionary arms race between adoption of camouflage to facilitate hunting and the ability of prey to quickly recognize predators by their formerly camouflaging patterns.

Time is a fundamental component of ecological processes. How animal behavior changes over time has been explored through well-known ecological theories like niche partitioning and predator-prey dynamics. Yet, changes in animal behavior within the shorter 24-hour light-dark cycle have largely gone unstudied. Understanding if an animal can adjust their temporal activity to mitigate or adapt to environmental change has become a recent topic of discussion and is important for effective wildlife management and conservation. While spatial habitat is a fundamental consideration in wildlife management and conservation, temporal habitat is often ignored. We formulated a temporal resource selection model to quantify the diel behavior of eight mammal species across ten U.S. cities. We found high variability in diel activity patterns within and among species and species-specific correlations between diel activity and human population density, impervious land cover, available greenspace, vegetation cover, and mean daily temperature. We also found that some species may modulate temporal behaviors to manage both natural and anthropogenic risks. Our results highlight the complexity with which temporal activity patterns interact with local environmental characteristics, and suggest that urban mammals may use time along the 24-hour cycle to reduce risk, adapt, and therefore persist, and in some cases thrive, in human-dominated ecosystems.

Primates are noted for their varied and complex pelage and bare skin coloration but the significance of this diverse coloration remains opaque. Using new updated information, novel scoring of coat and skin coloration, and controlling for shared ancestry, we reexamined and extended findings from previous studies across the whole order and the five major clades within it. Across primates, we found (i) direct and indirect evidence for pelage coloration being driven by protective coloration strategies including background matching, countershading, disruptive coloration, and aposematism, (ii) diurnal primates being more colorful, and (iii) the possibility that pelage color diversity is negatively associated with female trichromatic vision; while (iv) reaffirming avoidance of hybridization driving head coloration in males, (v) darker species living in warm, humid conditions (Gloger’s rule), and (vi) advertising to multiple mating partners favoring red genitalia in females. Nonetheless, the importance of these drivers varies greatly across clades. In strepsirrhines and cercopithecoids, countershading is important; greater color diversity may be important for conspecific signaling in more diurnal and social strepsirrhines; lack of female color vision may be associated with colorful strepsirrhines and platyrrhines; whereas cercopithecoids obey Gloger’s rule. Haplorrhines show background matching, aposematism, character displacement, and red female genitalia where several mating partners are available. Our findings emphasize several evolutionary drivers of coloration in this extraordinarily colorful order. Throughout, we used coarse but rigorous measures of coloration, and our ability to replicate findings from earlier studies opens up opportunities for classifying coloration of large numbers of species at a macroevolutionary scale.

Urban biodiversity provides critical ecosystem services and is a key component to environmentally and socially sustainable cities. However, biodiversity varies greatly within and among cities, leading to human communities with changing and unequal experiences with nature. The "luxury effect," a hypothesis that predicts a positive | 5447 MAGLE Et AL.

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