Thanatosis—also known as death-feigning and, we argue more appropriately, tonic immobility (TI)—is an under-reported but fascinating anti-predator strategy adopted by diverse prey late on in the predation sequence, and frequently following physical contact by the predator. TI is thought to inhibit further attack by predators and reduce the perceived need of the predator to subdue prey further. The behaviour is probably present in more taxa than is currently described, but even within well-studied groups the precise taxonomic distribution is unclear for a number of practical and ethical reasons. Here we synthesise the key studies investigating the form, function, evolutionary and ecological costs and benefits of TI. This review also considers the potential evolutionary influence of certain predator types in the development of the strategy in prey, and the other non-defensive contexts in which TI has been suggested to occur. We believe that there is a need for TI to be better appreciated in the scientific literature and outline potentially profitable avenues for investigation. Future use of technology in the wild should yield useful developments for this field of study.Significance statementAnti-predatory defences are crucial to many aspects of behavioural ecology. Thanatosis (often called death-feigning) has long been an under-appreciated defence, despite being taxonomically and ecologically widespread. We begin by providing much-needed clarification on both terminology and definition. We demonstrate how apparently disparate observations in the recent literature can be synthesised through placing the behaviour within a cost-benefit framework in comparison to alternative behavioural choices, and how aspects of the ecology differentially affect costs and benefits. Extending this, we provide novel insights into why the evolution of thanatosis can be understood in terms of coevolution between predators and prey. We offer further novel hypotheses, and discuss how these can be tested, focussing on how emerging technologies can be of great use in developing our understanding of thanatosis in free-living animals.
Dropping is a common antipredator defence that enables rapid escape from a perceived threat. However, despite its immediate effectiveness in predator–prey encounters (and against other dangers such as a parasitoid or an aggressive conspecific), it remains an under‐appreciated defence strategy in the scientific literature. Dropping has been recorded in a wide range of taxa, from primates to lizards, but has been studied most commonly in insects. Insects have been found to utilise dropping in response to both biotic and abiotic stimuli, sometimes dependent on mechanical or chemical cues. Whatever the trigger for dropping, the decision to drop by prey will present a range of inter‐related costs and benefits to the individual and so there will be subtle complexities in the trade‐offs surrounding this defensive behaviour. In predatory encounters, dropping by prey will also impose varying costs and benefits on the predator – or predators – involved in the system. There may be important trade‐offs involved in the decision made by predators regarding whether to pursue prey or not, but the predator perspective on dropping has been less explored at present. Beyond its function as an escape tactic, dropping has also been suggested to be an important precursor to flight in insects and further study could greatly improve understanding of its evolutionary importance. Dropping in insects could also prove of significant practical importance if an improved understanding can be applied to integrated pest‐management strategies. Currently the non‐consumptive effects of predators on their prey are under‐appreciated in biological control and it may be that the dropping behaviour of many pest species could be exploited via management practices to improve crop protection. Overall, this review aims to provide a comprehensive synthesis of the current literature on dropping and to raise awareness of this fascinating and widespread behaviour. It also seeks to offer some novel hypotheses and highlight key avenues for future research.
Deflection occurs in predator-prey interactions where prey possess traits that influence the position of the predator's initial contact with the prey's body in a way that enhances the prey's probability of survival when attacked. As an anti-predatory defence occurring late in the sequence of an attack, deflection is an understudied but fascinating strategy involving a range of unusual adaptations in diverse prey species. Deflective traits have been postulated to be important to the defensive strategies of a range of organisms, but while evidence for its existence is quite variable among groups, we argue that previous research neglects some promising taxa. As a defence, deflection will probably play a crucial role in the behavioural ecology and evolution of both prey species and their predators; as such it warrants greater interest from zoologists. Here, we first summarise what is known about deflection from the current literature. We next offer predictions about the coevolutionary possibilities surrounding deflection, based on the benefits and costs experienced by prey and their predators. Finally, we outline the most interesting outstanding avenues for future research in the field of deflection and make novel suggestions as to how they could be usefully explored.
Distraction displays are conspicuous behaviours functioning to distract a predator's attention away from the displayer's nest or young, thereby reducing the chance of offspring being discovered and predated. Distraction is one of the riskier parental care tactics, as its success derives from the displaying parent becoming the focus of a predator's attention. Such displays are prominent in birds, primarily shorebirds, but the last comprehensive review of distraction was in 1984. Our review aims to provide an updated synthesis of what is known about distraction displays in birds, and to open up new areas of study by highlighting some of the key avenues to explore and the broadened ecological perspectives that could be adopted in future research. We begin by drawing attention to the flexibility of form that distraction displays can take and providing an overview of the different avian taxa known to use anti‐predator distraction displays, also examining species‐specific sex differences in use. We then explore the adaptive value and evolution of distraction displays, before considering the variation seen in the timing of their use over a reproductive cycle. An evaluation of the efficacy of distraction compared with alternative anti‐predator tactics is then conducted via a cost–benefit analysis. Distraction displays are also found in a handful of non‐avian taxa, and we briefly consider these unusual cases. We conclude by postulating why distraction is primarily an avian behaviour and set out our suggestions for future research into the evolution and ecology of avian distraction displays.
Pearson's product moment correlation coefficient (more commonly Pearson's r) tends to underestimate correlations that exist in the underlying population. This phenomenon is generally unappreciated in studies of ecology, although a range of corrections are suggested in the statistical literature. The use of Pearson's r as the classical measure for correlation is widespread in ecology, where manipulative experiments are impractical across the large spatial scales concerned; it is therefore vital that ecologists are able to use this correlation measure as effectively as possible. Here, our literature review suggests that corrections for the issue of underestimation in Pearson's r should not be adopted if either the data deviate from bivariate normality or sample size is greater than around 30. Through our simulations, we then aim to offer advice to researchers in ecology on situations where both distributions can be described as normal, but sample sizes are lower than around 30. We found that none of the methods currently offered in the literature to correct the underestimation bias offer consistently reliable performance, and so we do not recommend that they be implemented when making inferences about the behaviour of a population from a sample. We also suggest that, when considering the importance of the bias towards underestimation in Pearson's product moment correlation coefficient for biological conclusions, the likely extent of the bias should be discussed. Unless sample size is very small, the issue of sample bias is unlikely to call for substantial modification of study conclusions.
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