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      Defense against predators incurs high reproductive costs for the aposematic moth Arctia plantaginis

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          Abstract

          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 ( h 2 = 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.

          Abstract

          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.

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          Most cited references48

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          Predator mixes and the conspicuousness of aposematic signals.

          Conspicuous warning signals of unprofitable prey are a defense against visually hunting predators. They work because predators learn to associate unprofitability with bright coloration and because strong signals are detectable and memorable. However, many species that can be considered defended are not very conspicuous; they have weak warning signals. This phenomenon has previously been ignored in models and experiments. In addition, there is significant within- and among-species variation among predators in their search behavior, in their visual, cognitive, and learning abilities, and in their resistance to defenses. In this article we explore the effects of variable predators on models that combine positive frequency-dependent, frequency-independent, and negative frequency-dependent predation and show that weak signaling of aposematic species can evolve if predators vary in their tendency to attack defended prey.
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            Why are distasteful prey not cryptic?

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              Co-mimics have a mutualistic relationship despite unequal defences.

              In the first clear mathematical treatment of natural selection, Müller proposed that a shared warning signal (mimicry) would benefit defended prey species by sharing out the per capita mortality incurred during predator education. Although mimicry is a mainstay of adaptationist thinking, there has been repeated debate on whether there is a mutualistic or a parasitic relationship between unequally defended co-mimic species. Here we show that the relationship between unequally defended species is mutualistic. We examined this in a 'novel world' of artificial prey with wild predators (great tit, Parus major). We kept the abundance of a highly defended prey ('model') constant and increased the density of a moderately defended prey ('defended mimic') of either perfect or imperfect mimetic resemblance to the model. Both model and defended mimic showed a net benefit from a density-dependent decrease in their per capita mortality. Even when the effect of dilution through density was controlled for, defended mimics did not induce additional attacks on the model, but we found selection for accurate signal mimicry. In comparison, the addition of fully edible (batesian) mimics did increase additional attacks on the model, but as a result of dilution this resulted in no overall increase in per capita mortality. By ignoring the effects of density, current theories may have overestimated the parasitic costs imposed by less defended mimics on highly defended models.

                Author and article information

                Contributors
                Role: Handling Editor
                Journal
                Behav Ecol
                Behav. Ecol
                beheco
                Behavioral Ecology
                Oxford University Press (UK )
                1045-2249
                1465-7279
                May-Jun 2020
                15 April 2020
                15 April 2020
                : 31
                : 3
                : 844-850
                Affiliations
                [1 ] Department of Biological and Environmental Sciences, University of Jyväskylä , Jyväskylä, Finland
                [2 ] Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam , Amsterdam, The Netherlands
                Author notes
                 Address correspondence to C. Lindstedt. E-mail: carita.a.lindstedt@ 123456jyu.fi

                These authors contributed equally to this study.

                Author information
                http://orcid.org/0000-0001-8176-3613
                http://orcid.org/0000-0003-2840-6267
                http://orcid.org/0000-0002-1117-5629
                Article
                araa033
                10.1093/beheco/araa033
                7303824
                194080b5-e79d-4c51-9119-1c8bf3a735b5
                © The Author(s) 2020. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 30 April 2019
                : 11 March 2020
                : 14 March 2020
                : 06 April 2020
                Page count
                Pages: 7
                Funding
                Funded by: Academy of Finland, DOI 10.13039/501100002341;
                Award ID: 136387
                Award ID: 257581
                Categories
                Original Articles
                AcademicSubjects/SCI01330

                Ecology
                chemical defense,color polymorphism,cost of defense,heritability
                Ecology
                chemical defense, color polymorphism, cost of defense, heritability

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