{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T06:40:47Z","timestamp":1790145647967,"version":"4.0.1"},"reference-count":104,"publisher":"PeerJ","license":[{"start":{"date-parts":[[2021,6,18]],"date-time":"2021-06-18T00:00:00Z","timestamp":1623974400000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100002341","name":"Academy of Finland","doi-asserted-by":"crossref","award":["286814"],"award-info":[{"award-number":["286814"]}],"id":[{"id":"10.13039\/501100002341","id-type":"DOI","asserted-by":"crossref"}]},{"name":"ERC","award":["339873"],"award-info":[{"award-number":["339873"]}]},{"name":"NERC Doctoral Training Partnership","award":["NE\/L002507\/1"],"award-info":[{"award-number":["NE\/L002507\/1"]}]},{"DOI":"10.13039\/501100004892","name":"Spanish Agency for International Development Cooperation","doi-asserted-by":"crossref","award":["2018SPE0000400194"],"award-info":[{"award-number":["2018SPE0000400194"]}],"id":[{"id":"10.13039\/501100004892","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>\n                    Chemical defences against predators underlie the evolution of aposematic coloration and mimicry, which are classic examples of adaptive evolution. Surprisingly little is known about the roles of ecological and evolutionary processes maintaining defence variation, and how they may feedback to shape the evolutionary dynamics of species. Cyanogenic\n                    <jats:italic>Heliconius<\/jats:italic>\n                    butterflies exhibit diverse warning color patterns and mimicry, thus providing a useful framework for investigating these questions. We studied intraspecific variation in de novo biosynthesized cyanogenic toxicity and its potential ecological and evolutionary sources in wild populations of\n                    <jats:italic>Heliconius erato<\/jats:italic>\n                    along environmental gradients, in common-garden broods and with feeding treatments. Our results demonstrate substantial intraspecific variation, including detectable variation among broods reared in a common garden. The latter estimate suggests considerable evolutionary potential in this trait, although predicting the response to selection is likely complicated due to the observed skewed distribution of toxicity values and the signatures of maternal contributions to the inheritance of toxicity. Larval diet contributed little to toxicity variation. Furthermore, toxicity profiles were similar along steep rainfall and altitudinal gradients, providing little evidence for these factors explaining variation in biosynthesized toxicity in natural populations. In contrast, there were striking differences in the chemical profiles of\n                    <jats:italic>H. erato<\/jats:italic>\n                    from geographically distant populations, implying potential local adaptation in the acquisition mechanisms and levels of defensive compounds. The results highlight the extensive variation and potential for adaptive evolution in defense traits for aposematic and mimetic species, which may contribute to the high diversity often found in these systems.\n                  <\/jats:p>","DOI":"10.7717\/peerj.11523","type":"journal-article","created":{"date-parts":[[2021,6,18]],"date-time":"2021-06-18T05:17:51Z","timestamp":1623993471000},"page":"e11523","source":"Crossref","is-referenced-by-count":16,"title":["Evolutionary and ecological processes influencing chemical defense variation in an aposematic and mimetic\n                    <i>Heliconius<\/i>\n                    butterfly"],"prefix":"10.7717","volume":"9","author":[{"given":"Anniina L. K.","family":"Mattila","sequence":"first","affiliation":[{"name":"Research Centre for Ecological Change, Organismal and Evolutionary Biology Research Programme, University of Helsinki, Helsinki, Finland"},{"name":"Helsinki Life Science Institute, University of Helsinki, Helsinki, Finland"},{"name":"*Current affiliation: Finnish Museum of Natural History (LUOMUS), University of Helsinki, Helsinki,  Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chris D.","family":"Jiggins","sequence":"additional","affiliation":[{"name":"Department of Zoology, University of Cambridge, Cambridge, United Kingdom"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00d8ystein H.","family":"Opedal","sequence":"additional","affiliation":[{"name":"Department of Biology, Lund University, Lund, Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gabriela","family":"Montejo-Kovacevich","sequence":"additional","affiliation":[{"name":"Department of Zoology, University of Cambridge, Cambridge, United Kingdom"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00c9rika C.","family":"Pinheiro de castro","sequence":"additional","affiliation":[{"name":"Department of Zoology, University of Cambridge, Cambridge, United Kingdom"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"W. Owen","family":"McMillan","sequence":"additional","affiliation":[{"name":"Smithsonian Tropical Research Institute, Gamboa, Panama"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Caroline","family":"Bacquet","sequence":"additional","affiliation":[{"name":"Universidad Regional Amaz\u00f3nica de Ikiam, Tena, Ecuador"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marjo","family":"Saastamoinen","sequence":"additional","affiliation":[{"name":"Research Centre for Ecological Change, Organismal and Evolutionary Biology Research Programme, University of Helsinki, Helsinki, Finland"},{"name":"Helsinki Life Science Institute, University of Helsinki, Helsinki, Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4443","published-online":{"date-parts":[[2021,6,18]]},"reference":[{"key":"10.7717\/peerj.11523\/ref-1","doi-asserted-by":"publisher","first-page":"11021","DOI":"10.1038\/srep11021","article-title":"Signal honesty and predation risk among a closely related group of aposematic species","volume":"5","author":"Arenas","year":"2015","journal-title":"Scientific Reports"},{"key":"10.7717\/peerj.11523\/ref-2","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1186\/s12862-016-0843-5","article-title":"Variation in cyanogenic compounds concentration within a Heliconius butterfly community: does mimicry explain everything?","volume":"16","author":"Arias","year":"2016","journal-title":"BMC Evolutionary Biology"},{"key":"10.7717\/peerj.11523\/ref-3","doi-asserted-by":"publisher","first-page":"20140533","DOI":"10.1098\/rsbl.2014.0533","article-title":"Better the devil you know: avian predators find variation in prey toxicity aversive","volume":"10","author":"Barnett","year":"2014","journal-title":"Biology Letters"},{"key":"10.7717\/peerj.11523\/ref-4","article-title":"Fitting linear mixed-effects models using lme4","author":"Bates","year":"2014","unstructured":"arXiv preprint. arXiv:1406.5823"},{"key":"10.7717\/peerj.11523\/ref-5","doi-asserted-by":"publisher","first-page":"936","DOI":"10.1126\/science.176.4037.936","article-title":"Natural selection for M\u00fcllerian mimicry in Heliconius erato in Costa Rica","volume":"176","author":"Benson","year":"1972","journal-title":"Science"},{"key":"10.7717\/peerj.11523\/ref-6","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1111\/j.1558-5646.1975.tb00861.x","article-title":"Coevolution of plants and herbivores: passion flower butterflies","volume":"29","author":"Benson","year":"1975","journal-title":"Evolution"},{"key":"10.7717\/peerj.11523\/ref-7","doi-asserted-by":"publisher","first-page":"334","DOI":"10.1111\/j.1365-2435.2012.01961.x","article-title":"How the ladybird got its spots: effects of resource limitation on the honesty of aposematic signals","volume":"26","author":"Blount","year":"2012","journal-title":"Functional Ecology"},{"key":"10.7717\/peerj.11523\/ref-8","doi-asserted-by":"publisher","first-page":"871","DOI":"10.1098\/rspb.2008.1407","article-title":"Warning displays may function as honest signals of toxicity","volume":"276","author":"Blount","year":"2009","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"key":"10.7717\/peerj.11523\/ref-9","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1007\/s00442-005-0076-6","article-title":"Larval food limitation in butterflies: effects on adult resource allocation and fitness","volume":"144","author":"Boggs","year":"2005","journal-title":"Oecologia"},{"key":"10.7717\/peerj.11523\/ref-10","doi-asserted-by":"publisher","first-page":"2703","DOI":"10.1111\/evo.13368","article-title":"Selection on skewed characters and the paradox of stasis","volume":"71","author":"Bonamour","year":"2017","journal-title":"Evolution"},{"key":"10.7717\/peerj.11523\/ref-11","first-page":"216","article-title":"The evolution of unpalatability and the cost of chemical defense in insects","volume-title":"Insect chemical ecology: an evolutionary approach","author":"Bowers","year":"1992"},{"key":"10.7717\/peerj.11523\/ref-12","article-title":"R package sm: nonparametric smoothing methods","author":"Bowman","year":"2014"},{"key":"10.7717\/peerj.11523\/ref-13","doi-asserted-by":"publisher","first-page":"73","DOI":"10.2307\/2407673","article-title":"Butterfly palatability and mimicry: experiments with ameiva lizards","volume":"30","author":"Boyden","year":"1976","journal-title":"Evolution"},{"key":"10.7717\/peerj.11523\/ref-14","doi-asserted-by":"publisher","first-page":"388","DOI":"10.1111\/brv.12460","article-title":"Diversity in warning coloration: selective paradox or the norm?","volume":"94","author":"Briolat","year":"2019","journal-title":"Biological Reviews"},{"key":"10.7717\/peerj.11523\/ref-15","doi-asserted-by":"publisher","first-page":"1059","DOI":"10.1073\/pnas.66.4.1059","article-title":"Theoretical investigations of automimicry, I. 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