Spatiotemporal dynamics of animal contests arise from effective forces between contestants

Competition among animals for resources, notably food, territories, and mates, is ubiquitous at all scales of life. This competition is often resolved through contests among individuals, which are commonly understood according to their outcomes and in particular, how these outcomes depend on decision-making by the contestants. Because they are restricted to end-point predictions, these approaches cannot predict real-time or real-space dynamics of animal contest behavior. This limitation can be overcome by studying systems that feature typical contest behavior while being simple enough to track and model. Here, we propose to use such systems to construct a theoretical framework that describes real-time movements and behaviors of animal contestants. We study the spatiotemporal dynamics of contests in an orb-weaving spider, in which all the common elements of animal contests play out. The confined arena of the web, on which interactions are dominated by vibratory cues in a two-dimensional space, simplifies the analysis of interagent interactions. We ask whether these seemingly complex decision-makers can be modeled as interacting active particles responding only to effective forces of attraction and repulsion due to their interactions. By analyzing the emergent dynamics of “contestant particles,” we provide mechanistic explanations for real-time dynamical aspects of animal contests, thereby explaining competitive advantages of larger competitors and demonstrating that complex decision-making need not be invoked in animal contests to achieve adaptive outcomes. Our results demonstrate that physics-based classification and modeling, in terms of effective rules of interaction, provide a powerful framework for understanding animal contest behaviors.

Animal contests and microplastics: evidence of disrupted behaviour in hermit crabs Pagurus bernhardus

Microplastics are ubiquitous in global marine systems and may have negative impacts on a vast range of species. Recently, microplastics were shown to impair shell selection assessments in hermit crabs, an essential behaviour for their survival. Hermit crabs also engage in ‘rapping’ contests over shells, based on cognitive assessments of shell quality and opponent fighting ability and, hence, are a useful model species for examining the effects of microplastics on fitness-relevant behaviour in marine systems. Here, we investigated how a 5-day microplastic exposure (25 microplastics/litre) affected the dynamics and outcome of 120 staged hermit crab contests. Using a 2 × 2 factorial design, we examined how microplastics (i.e. presence or absence) and contestant role (i.e. attacker or defender) affected various behavioural variables. Significantly higher raps per bout were needed to evict microplastic-treated defenders when attackers were pre-exposed to control conditions (i.e. no plastic). Also, significantly longer durations of rapping bouts were needed to evict control-treated defenders when attackers were pre-exposed to microplastics. We suggest that microplastics impaired defenders' ability to identify resource holding potential and also affected attackers’ rapping strength and intensity during contests. These impacts on animal contests indicate that microplastics have broader deleterious effects on marine biota than currently recognized.

Repeatability of combat rate across different group compositions in male house finches

Studies of animal contests have focused on the probability of winning an encounter, because it directly affects the benefits of competition. However, the costs (e.g., physiological stress) and benefits of competition should also depend on the number of aggressive encounters per unit time (combat rate, hereafter) in which the focal individual is involved. Using colourful and drab male house finches (Haemorhous mexicanus) from urban and rural sites, we showed that combat rate was repeatable across the same and different group sizes for birds who won competitions. In addition, colourful urban males exhibited the lowest propensity for frequent aggression (and hence low combat rate). However, male bill size (another trait we previously found to correlate with male competitiveness in this species) was not related to aggressive propensity. Combat rate can be predicted by male identity and some, but not all, predictors of male competitiveness.

Using knowledge from human research to improve understanding of contest theory and contest dynamics

Our understanding of animal contests and the factors that affect contest dynamics and decisions stems from a long and prosperous collaboration between empiricists and theoreticians. Over the last two decades, however, theoretical predictions regarding the factors that affect individual decisions before, during and after a contest are becoming increasingly difficult to test empirically. Extremely large sample sizes are necessary to experimentally test the nuanced theoretical assumptions surrounding how information is used by animals during a contest, how context changes the information used, and how individuals change behaviour as a result of both the information available and the context in which the information is acquired. In this review, we discuss how the investigation of contests in humans through the collaboration of biologists and psychologists may advance contest theory and dynamics in general. We argue that a long and productive history exploring human behaviour and psychology combined with technological advancements provide a unique opportunity to manipulate human perception during contests and collect unbiased data, allowing more targeted examinations of particular aspects of contest theory (e.g. winner/loser effects, information use as a function of age). We hope that our perspective provides the impetus for many future collaborations between biologists and psychologists.

The role of skill in animal contests: a neglected component of fighting ability

What attributes make some individuals more likely to win a fight than others? A range of morphological and physiological traits have been studied intensely but far less focus has been placed on the actual agonistic behaviours used. Current studies of agonistic behaviour focus on contest duration and the vigour of fighting. It also seems obvious that individuals that fight more skilfully should have a greater chance of winning a fight. Here, we discuss the meaning of skill in animal fights. As the activities of each opponent can be disrupted by the behaviour of their rival, we differentiate among ability, technique and skill itself. In addition to efficient, accurate and sometimes precise movement, skilful fighting also requires rapid decision-making, so that appropriate tactics and strategies are selected. We consider how these different components of skill could be acquired, through genes, experiences of play-fighting and of real fights. Skilful fighting can enhance resource holding potential (RHP) by allowing for sustained vigour, by inflicting greater costs on opponents and by minimizing the chance of damage. Therefore, we argue that skill is a neglected but important component of RHP that could be readily studied to provide new insights into the evolution of agonistic behaviour.