It is not always about body size: evidence of Rensch's rule in a male weapon.

In many species, sexual dimorphism increases with body size when males are the larger sex but decreases when females are the larger sex, a macro-evolutionary pattern known as Rensch's rule (RR). Although empirical studies usually focus exclusively on body size, Rensch's original proposal included sexual differences in other traits, such as ornaments and weapons. Here, we used a clade of harvestmen to investigate whether two traits follow RR: body size and length of the fourth pair of legs (legs IV), which are used as weapons in male-male fights. We found that males were slightly smaller than females and body size did not follow RR, whereas legs IV were much longer in males and followed RR. We propose that sexual selection might be stronger on legs IV length than on body size in males, and we discuss the potential role of condition dependence in the emergence of RR.

Flight initiation distance, color and camouflage.

Camouflage is widespread throughout the animal kingdom allowing individuals to avoid detection and hence save time and energy rather than escape from an approaching predator. Thus, camouflage is likely to have co-evolved with antipredator behavior. Here, we propose that camouflage results in dichotomous escape behavior within and among species with classes of individuals and species with cryptic coloration having shorter flight initiation distances (FIDs; the distance at which an individual takes flight when approached by a human). We report the results of 2 tests of this hypothesis. First, bird species with cryptically colored plumage have consistently shorter FID than closely related species without such color. Within species with sexually dimorphic plumage, brightly colored adult male common pheasants Phasianus colchicus and golden pheasants Chrysolophus pictus have long and variable FID, whereas cryptically colored juveniles and adult females have short and invariable FID. Second, FID in females was predicted by presence or absence of cryptic color, FID in males and their interaction. These findings are consistent with the hypothesis that risk-taking behavior has been attuned to camouflage, and that species with different levels of camouflage differ consistently in their FID.

Fifty years of chasing lizards: new insights advance optimal escape theory.

Systematic reviews and meta-analyses often examine data from diverse taxa to identify general patterns of effect sizes. Meta-analyses that focus on identifying generalisations in a single taxon are also valuable because species in a taxon are more likely to share similar unique constraints. We conducted a comprehensive phylogenetic meta-analysis of flight initiation distance in lizards. Flight initiation distance (FID) is a common metric used to quantify risk-taking and has previously been shown to reflect adaptive decision-making. The past decade has seen an explosion of studies focused on quantifying FID in lizards, and, because lizards occur in a wide range of habitats, are ecologically diverse, and are typically smaller and differ physiologically from the better studied mammals and birds, they are worthy of detailed examination. We found that variables that reflect the costs or benefits of flight (being engaged in social interactions, having food available) as well as certain predator effects (predator size and approach speed) had large effects on FID in the directions predicted by optimal escape theory. Variables that were associated with morphology (with the exception of crypsis) and physiology had relatively small effects, whereas habitat selection factors typically had moderate to large effect sizes. Lizards, like other taxa, are very sensitive to the costs of flight.

Increased tolerance to humans among disturbed wildlife.

Human disturbance drives the decline of many species, both directly and indirectly. Nonetheless, some species do particularly well around humans. One mechanism that may explain coexistence is the degree to which a species tolerates human disturbance. Here we provide a comprehensive meta-analysis of birds, mammals and lizards to investigate species tolerance of human disturbance and explore the drivers of this tolerance in birds. We find that, overall, disturbed populations of the three major taxa are more tolerant of human disturbance than less disturbed populations. The best predictors of the direction and magnitude of bird tolerance of human disturbance are the type of disturbed area (urbanized birds are more tolerant than rural or suburban populations) and body mass (large birds are more tolerant than small birds). By identifying specific features associated with tolerance, these results guide evidence-based conservation strategies to predict and manage the impacts of increasing human disturbance on birds.

How Nature-Based Tourism Might Increase Prey Vulnerability to Predators.

Tourism can be deleterious for wildlife because it triggers behavioral changes in individuals with cascading effects on populations and communities. Among these behavioral changes, animals around humans often reduce their fearfulness and antipredator responses towards humans. A straightforward prediction is that habituation to humans associated with tourism would negatively influence reaction to predators. This could happen indirectly, where human presence decreases the number of natural predators and thus prey become less wary, or directly, where human-habituated individuals become bolder and thus more vulnerable to predation. Building on ideas from the study of traits associated with domestication and urbanization, we develop a framework to understand how behavioral changes associated with nature-based tourism can impact individual fitness, and thus the demographic trajectory of a population.

Brain size as a driver of avian escape strategy.

After detecting an approaching predator, animals make a decision when to flee. Prey will initiate flight soon after detecting a predator so as to minimize attentional costs related to on-going monitoring of the whereabouts of the predator. Such costs may compete with foraging and other maintenance activities and hence be larger than the costs of immediate flight. The drivers of interspecific variation in escape strategy are poorly known. Here we investigated the morphological, life history and natural history traits that correlate with variation in avian escape strategy across a sample of 96 species of birds. Brain mass, body size, habitat structure and group size were the main predictors of escape strategy. The direction of the effect of these traits was consistent with selection for a reduction of monitoring costs. Therefore, attentional costs depend on relative brain size, which determines the ability to monitor the whereabouts of potential predators and the difficulty of this task as reflected by habitat and social complexity. Thus brain size, and the cognitive functions associated with it, constitute a general framework for explaining the effects of body size, habitat structure and sociality identified as determinants of avian escape strategy.

Birds flush early and avoid the rush: an interspecific study.

Since 1986, studies about the escape decisions made by prey are grounded in optimal escape theory (OET) which states that prey will initiate escape when the risk of remaining and the costs of leaving are equal. However, a recent hypothesis, Flush Early and Avoid the Rush (FEAR), acknowledged that the cost of monitoring approaching predators might be a ubiquitous cost. The FEAR hypothesis predicts that prey will generally flee soon after they detect a predator so as to minimize the costs incurred by monitoring the predator. Knowing whether animals flee to reduce monitoring costs is of applied interest because wildlife managers use escape behavior to create set-back zones to reduce human-wildlife conflict. Here we provide the most comprehensive assessment of the FEAR hypothesis using data collected from 178 bird species representing 67 families from two continents. The FEAR hypothesis explains escape behavior in 79% of studied species. Because the FEAR hypothesis is a widespread phenomenon that drives escape behavior in birds, alert distance must be systematically incorporated into the design of set-back zones to protect vulnerable species.

Sex differences in lizard escape decisions vary with latitude, but not sexual dimorphism.

Sexual selection is a powerful evolutionary mechanism that has shaped the physiology, behaviour and morphology of the sexes to the extent that it can reduce viability while promoting traits that enhance reproductive success. Predation is one of the underlying mechanisms accounting for viability costs of sexual displays. Therefore, we should expect that individuals of the two sexes adjust their anti-predator behaviour in response to changes in predation risk. We conducted a meta-analysis of 28 studies (42 species) of sex differences in risk-taking behaviour in lizards and tested whether these differences could be explained by sexual dichromatism, by sexual size dimorphism or by latitude. Latitude was the best predictor of the interspecific heterogeneity in sex-specific behaviour. Males did not change their escape behaviour with latitude, whereas females had increasingly reduced wariness at higher latitudes. We hypothesize that this sex difference in risk-taking behaviour is linked to sex-specific environmental constraints that more strongly affect the reproductive effort of females than males. This novel latitudinal effect on sex-specific anti-predator behaviour has important implications for responses to climate change and for the relative roles of natural and sexual selection in different species.

Phi index: a new metric to test the flush early and avoid the rush hypothesis.

Optimal escape theory states that animals should counterbalance the costs and benefits of flight when escaping from a potential predator. However, in apparent contradiction with this well-established optimality model, birds and mammals generally initiate escape soon after beginning to monitor an approaching threat, a phenomena codified as the "Flush Early and Avoid the Rush" (FEAR) hypothesis. Typically, the FEAR hypothesis is tested using correlational statistics and is supported when there is a strong relationship between the distance at which an individual first responds behaviorally to an approaching predator (alert distance, AD), and its flight initiation distance (the distance at which it flees the approaching predator, FID). However, such correlational statistics are both inadequate to analyze relationships constrained by an envelope (such as that in the AD-FID relationship) and are sensitive to outliers with high leverage, which can lead one to erroneous conclusions. To overcome these statistical concerns we develop the phi index (Φ), a distribution-free metric to evaluate the goodness of fit of a 1:1 relationship in a constraint envelope (the prediction of the FEAR hypothesis). Using both simulation and empirical data, we conclude that Φ is superior to traditional correlational analyses because it explicitly tests the FEAR prediction, is robust to outliers, and it controls for the disproportionate influence of observations from large predictor values (caused by the constrained envelope in AD-FID relationship). Importantly, by analyzing the empirical data we corroborate the strong effect that alertness has on flight as stated by the FEAR hypothesis.

American exceptionalism: population trends and flight initiation distances in birds from three continents.

BACKGROUND: All organisms may be affected by humans' increasing impact on Earth, but there are many potential drivers of population trends and the relative importance of each remains largely unknown. The causes of spatial patterns in population trends and their relationship with animal responses to human proximity are even less known. METHODOLOGY/PRINCIPAL FINDING: We investigated the relationship between population trends of 193 species of bird in North America, Australia and Europe and flight initiation distance (FID); the distance at which birds take flight when approached by a human. While there is an expected negative relationship between population trend and FID in Australia and Europe, we found the inverse relationship for North American birds; thus FID cannot be used as a universal predictor of vulnerability of birds. However, the analysis of the joint explanatory ability of multiple drivers (farmland breeding habitat, pole-most breeding latitude, migratory habit, FID) effects on population status replicated previously reported strong effects of farmland breeding habitat (an effect apparently driven mostly by European birds), as well as strong effects of FID, body size, migratory habit and continent. Farmland birds are generally declining. CONCLUSIONS/SIGNIFICANCE: Flight initiation distance is related to population trends in a way that differs among continents opening new research possibilities concerning the causes of geographic differences in patterns of anti-predator behavior.

Do animals generally flush early and avoid the rush? A meta-analysis.

Optimal escape theory predicts that animals should balance the costs and benefits of flight. One cost of not fleeing is the ongoing cost of monitoring an approaching predator. We used a phylogenetic meta-analysis to test the general hypothesis that animals should initiate flight soon after they detect a predator-the 'flush early and avoid the rush' hypothesis. We found a large, significant overall relationship between the distance at which animals were approached or first detected a threat and the distance at which they fled. While these results are the first general test of the flush early and avoid the rush hypothesis, future work will be required to determine whether animals flush early to reduce ongoing attentional costs, or if they flush early as a form of risk reduction.