Turn alternation and the influence of environmental factors on search routes through branched structures by ladybirds (Coccinella septempunctata and Adalia bipunctata).

Turn alternation is a locomotory behaviour wherein an animal makes consecutive turns in opposite directions (left-then-right or right-then-left). It has been suggested that its adaptive function is to maintain locomotion in a relatively constant general direction while negotiating obstacles. Previous work has focussed on the use of turn alternation in prey species in artificial horizontal mazes. In the first study presented here, we tested whether predatory seven-spot ladybirds (Coccinella septempunctata) exhibit turn alternation when repeatedly presented with consecutive choice turning decisions on vertically oriented twigs. Our findings suggest that turn alternation occurs vertically as well as horizontally, on plant structures as well as terrestrially, and in a type of animal (predatory insect) in which turn alternation has received little attention. In the second study presented here, we tested whether characteristics of branched structures explored by two-spot ladybirds (Adalia bipunctata) influence turning decisions. Our findings suggest that ladybirds exhibit preferences for thicker over thinner, straighter over more deviating headings, and higher over lower turning choice options at bifurcations. These exploratory studies indicate that while turn alternation is an observable phenomenon in ladybirds, it is not the only predictor of searching behaviour on branched structures.

The dicey dinner dilemma: Asymmetry in predator-prey risk-taking, a broadly applicable alternative to the life-dinner principle.

Forty years ago, the 'life-dinner principle' was proposed as an example of an asymmetry that may lead prey species to experience stronger selection than their predators, thus accounting for the high frequency with which prey escape alive from interaction with a predator. This principle remains an influential concept in the scientific literature, despite several works suggesting that the concept relies on many under-appreciated assumptions and does not apply as generally as was initially proposed. Here, we present a novel model describing a very different asymmetry to that proposed in the life-dinner principle, but one that could apply broadly. We argue that asymmetries between the relative costs and benefits to predators and prey of selecting a risky behaviour during an extended predator-prey encounter could lead to an enhanced likelihood of escape for the prey. Any resulting advantage to prey depends upon there being a behaviour or choice that introduces some inherent danger to both predator and prey if they adopt it, but which if the prey adopts the predator must match in order to have a chance of successful predation. We suggest that the circumstances indicated by our model could apply broadly across diverse taxa, including both risky spatial or behavioural choices.

Adaptive suicide: is a kin-selected driver of fatal behaviours likely?

While several manipulated host behaviours are accepted as extended phenotypes of parasites, there remains debate over whether other altered behaviours in hosts following parasitic invasion represent cases of parasite manipulation, host defence or the pathology of infection. One particularly controversial subject is 'suicidal behaviour' in infected hosts. The host-suicide hypothesis proposes that host death benefits hosts doomed to reduced direct fitness by protecting kin from parasitism and therefore increasing inclusive fitness. However, adaptive suicide has been difficult to demonstrate conclusively as a host adaptation in studies on social or clonal insects, for whom high relatedness should enable greater inclusive fitness benefits. Following discussion of empirical and theoretical works from a behavioural ecology perspective, this review finds that the most persuasive evidence for selection of adaptive suicide comes from bacteria. Despite a focus on parasites, driven by the existing literature, the potential for the evolution of adaptive suicidal behaviour in hosts is also considered to apply to cases of infection by pathogens, provided that the disease has a severe effect on direct fitness and that suicidal behaviour can affect pathogen transmission dynamics. Suggestions are made for future research and a broadening of the possible implications for coevolution between parasites and hosts.

Dropping to escape: a review of an under-appreciated antipredator defence.

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.

Underestimation of Pearson's product moment correlation statistic.

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.

A review of thanatosis (death feigning) as an anti-predator behaviour.

ABSTRACT: 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 STATEMENT: Anti-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.

Consequences of grouped data for testing for departure from circular uniformity.

Limits to the precision of circular data often cause grouping of data points into discrete categories, but the effects of grouping on tests for circular uniformity have been little explored. The Rayleigh test is often applied to grouped circular data, despite it being designed for continuous data and the statistical literature recommending a suite of alternative tests specifically designed for grouped data. Here, we investigated the performance of the Rayleigh test relative to four alternatives for testing the null hypothesis of uniformity in grouped circular data. We employed simulation, grouping data into a discrete number of same-sized categories and with samples drawn from a range of different distributions. We found that grouping had little effect on the type I error rate or the power of the Rayleigh test, and that the power of the Rayleigh test was very similar to that of the previously recommended alternative tests designed specifically for grouped circular data. It may thus be appropriate to apply the Rayleigh test to grouped data, provided the situation is one in which the test has substantial statistical power.