Prey seldom rely on a single type of antipredator defence, often using multiple defences to avoid predation. In many cases, selection in different contexts may favour the evolution of multiple defences in a prey. However, a prey may use multiple defences to protect itself during a single predator encounter. Such "defence portfolios" that defend prey against a single instance of predation are distributed across and within successive stages of the predation sequence (encounter, detection, identification, approach (attack), subjugation and consumption). We contend that at present, our understanding of defence portfolio evolution is incomplete, and seen from the fragmentary perspective of specific sensory systems (e.g., visual) or specific types of defences (especially aposematism). In this review, we aim to build a comprehensive framework for conceptualizing the evolution of multiple prey defences, beginning with hypotheses for the evolution of multiple defences in general, and defence portfolios in particular. We then examine idealized models of resource trade-offs and functional interactions between traits, along with evidence supporting them. We find that defence portfolios are constrained by resource allocation to other aspects of life history, as well as functional incompatibilities between different defences. We also find that selection is likely to favour combinations of defences that have synergistic effects on predator behaviour and prey survival. Next, we examine specific aspects of prey ecology, genetics and development, and predator cognition that modify the predictions of current hypotheses or introduce competing hypotheses. We outline schema for gathering data on the distribution of prey defences across species and geography, determining how multiple defences are produced, and testing the proximate mechanisms by which multiple prey defences impact predator behaviour. Adopting these approaches will strengthen our understanding of multiple defensive strategies.

CEFE Université de Montpellier CNRS EPHE IRD Montpellier France

Centre for Ecology and Conservation University of Exeter Penryn UK

Computer Vision Center Computer Science Department Universitat Autònoma de Barcelona Barcelona Spain

Department of Biological Sciences California State University Long Beach California USA

Department of Biological Sciences Purdue University West Lafayette Indiana USA

Department of Biology and Environmental Science University of Jyväskylä Jyväskylä Finland

Department of Biology Carleton University Ottawa Ontario Canada

Department of Biology East Carolina University Greenville North Carolina USA

Department of Biology Trent University Peterborough Ontario Canada

Department of Biology University of North Carolina at Chapel Hill Chapel Hill North Carolina USA

Department of Biosciences Swansea University Swansea UK

Department of Botany and Zoology Faculty of Science Masaryk University Brno Czech Republic

Department of Collective Behavior Max Planck Institute of Animal Behavior Konstanz Germany

Department of Forest Sciences University of Helsinki Helsinki Finland

Department of Integrative Biology Oregon State University Corvallis Oregon USA

Department of Interdisciplinary Life Sciences Konrad Lorenz Institute of Ethology University of Veterinary Medicine Vienna Austria

Department of Zoology Faculty of Science Charles University Prague Czech Republic

Evolutionary Biology Universität Bielefeld Bielefeld Germany

Grupo de Biodiversidad Medio Ambiente y Salud Universidad de Las Américas Quito Ecuador

Hawkesbury Institute of the Environment Western Sydney University Penrith New South Wales Australia

HiLIFE Helsinki Institute of Life Sciences University of Helsinki Helsinki Finland

Institut de Systématique Evolution Biodiversité CNRS MNHN Sorbonne Université EPHE Université des Antilles Paris France

Institute of Biodiversity and Ecosystem Dynamics University of Amsterdam Amsterdam The Netherlands

Institute of Biosciences Faculty of Medical Sciences Newcastle University Newcastle upon Tyne UK

Institute of Integrative Biology ETH Zurich Zurich Switzerland

Laboratoire Écologie Évolution Interactions des Systèmes Amazoniens Université de Guyane CNRS IFREMER Cayenne France

Laboratoire Évolution and Diversité Biologique Université Paul Sabatier Toulouse 3 UMR 5174 CNRS IRD Toulouse France

Max Planck Research Group Predators and Toxic Prey Max Planck Institute for Chemical Ecology Jena Germany

National Centre for Biological Sciences Tata Institute of Fundamental Research Bengaluru India

Research Programme in Organismal and Evolutionary Biology Faculty of Biological and Environmental Sciences University of Helsinki Helsinki Finland

School of Biological Sciences The University of Queensland St Lucia Queensland Australia

School of Biological Sciences University of Bristol Bristol UK

School of BioSciences University of Melbourne Melbourne Victoria Australia

School of Life Sciences University of Nottingham Nottingham UK

School of Natural Sciences Macquarie University Sydney New South Wales Australia

School of Psychology and Neuroscience University of St Andrews St Andrews UK

School of Science Western Sydney University Penrith New South Wales Australia

Smithsonian Tropical Research Institute Gamboa Panama

Zukunftskolleg University of Konstanz Konstanz Germany

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I remember you! Multicomponent warning signals and predator memory

Quantifying the success of prey crypsis, aposematism, and evasiveness in avoiding predator attack