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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. peerReviewed

Additional file 1: Pilot hexane extractions.

Systematic literature review conducted to answer the question "Does a standardised Morbidity and Mortality meeting (M&M) structure and process support learning and system improvement and improve clinician engagement and patient safety culture?" Inclusion and exclusion criteria Empirical or theoretical studies discussing M&Ms in English focused on adult populations and hospital settings were included. Publications that were reviews or opinion pieces were excluded along with abstracts or conference presentation summaries. Studies measuring and reporting clinical rather than process outcomes were excluded as not relevant to the purpose of this review. Electronic databases A range of text words, synonyms, and subject headings were developed for M&Ms (see Excel file). These text words, synonyms, and subject headings were used to search three electronic databases that index journals of relevance to the review topic (Pubmed, Embase and CINAHL).

To understand how variation in warning displays evolves and is maintained, we need to understand not only how perceivers of these traits select color and toxicity but also the sources of the genetic and phenotypic variation exposed to selection by them. We studied these aspects in the wood tiger moth Arctia plantaginis, which has two locally co-occurring male color morphs in Europe: yellow and white. When threatened, both morphs produce defensive secretions from their abdomen and from thoracic glands. Abdominal fluid has shown to be more important against invertebrate predators than avian predators, and the defensive secretion of the yellow morph is more effective against ants. Here, we focused on the morph-linked reproductive costs of secretion of the abdominal fluid and quantified the proportion of phenotypic and genetic variation in it. We hypothesized that, if yellow males pay higher reproductive costs for their more effective aposematic display, the subsequent higher mating success of white males could offer one explanation for the maintenance of the polymorphism. We first found that the heritable variation in the quantity of abdominal secretion was very low (h2 = 0.006) and the quantity of defensive secretion was not dependent on the male morph. Second, deploying the abdominal defensive secretion decreased the reproductive output of both color morphs equally. This suggests that potential costs of pigment production and chemical defense against invertebrates are not linked in A. plantaginis. Furthermore, our results indicate that environmentally induced variation in chemical defense can alter an individual’s fitness significantly., How much does it cost to defend against predators? A lot if you are a male wood tiger moth as deploying their defensive secretion decreases their reproductive success. When threatened, these aposematic moths produce defensive secretions from their abdomen, which are targeted specifically against invertebrate predators. However, this invertebrate-specific defense does not come without costs: deploying the defensive secretion only once decreases the reproductive success of males.