{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,2,21]],"date-time":"2025-02-21T03:43:45Z","timestamp":1740109425177,"version":"3.37.3"},"reference-count":52,"publisher":"Oxford University Press (OUP)","issue":"7","license":[{"start":{"date-parts":[[2023,7,1]],"date-time":"2023-07-01T00:00:00Z","timestamp":1688169600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"},{"start":{"date-parts":[[2023,2,3]],"date-time":"2023-02-03T00:00:00Z","timestamp":1675382400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"funder":[{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["NRF- 2019R1C1C1002466"],"award-info":[{"award-number":["NRF- 2019R1C1C1002466"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["NRF\u2010 2019R1C1C1002466"],"award-info":[{"award-number":["NRF\u2010 2019R1C1C1002466"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2023,7,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Countershading is a gradient of colouration in which the illuminated dorsal surfaces are darker than the unilluminated ventral surface. It is widespread in the animal kingdom and endows the body with a more uniform colour to decrease the chance of detection by predators. Although recent empirical studies support the theory of survival advantage conferred by countershading, this camouflage strategy has evolved only in some of the cryptic animals, and our understanding of the factors that affect the evolution of countershading is limited. This study examined the association between body size and countershading using lepidopteran larvae (caterpillars) as a model system. Specifically, we predicted that countershading may have selectively evolved in large-sized species among cryptic caterpillars if (1) large size constrains camouflage which facilitates the evolution of a trait reinforcing their crypsis and (2) the survival advantage of countershading is size-dependent. Phylogenetic analyses of four different lepidopteran families (Saturniidae, Sphingidae, Erebidae, and Geometridae) suggest equivocal results: countershading was more likely to be found in larger species in Saturniidae but not in the other families. The field predation experiment assuming avian predators did not support size-dependent predation in countershaded prey. Collectively, we found only weak evidence that body size is associated with countershading in caterpillars. Our results suggest that body size is not a universal factor that has shaped the interspecific variation in countershading observed in caterpillars.<\/jats:p>\n               <jats:p>Abstract<\/jats:p>\n               <jats:p>We tested whether there exists an association between body size and countershading in caterpillars using phylogenetic analyses and field predation experiments. We found only weak evidence that body size is associated with countershading in caterpillars.<\/jats:p>","DOI":"10.1111\/jeb.14153","type":"journal-article","created":{"date-parts":[[2023,2,4]],"date-time":"2023-02-04T04:43:58Z","timestamp":1675485838000},"page":"1032-1039","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Comparative and experimental studies on the relationship between body size and countershading in caterpillars"],"prefix":"10.1093","volume":"36","author":[{"given":"Yerin","family":"Hwang","sequence":"first","affiliation":[{"name":"Department of Agricultural Biotechnology Seoul National University Seoul South Korea"},{"name":"Department of Biosciences Mokpo National University Muan South Korea"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sohee","family":"Yoo","sequence":"additional","affiliation":[{"name":"Department of Agricultural Biotechnology Seoul National University Seoul South Korea"},{"name":"Department of Biosciences Mokpo National University Muan South Korea"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Chohee","family":"Park","sequence":"additional","affiliation":[{"name":"Department of Agricultural Biotechnology Seoul National University Seoul South Korea"},{"name":"Department of Biosciences Mokpo National University Muan South Korea"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3707-4989","authenticated-orcid":false,"given":"Changku","family":"Kang","sequence":"additional","affiliation":[{"name":"Research Institute of Agriculture and Life Sciences Seoul National University Seoul South Korea"},{"name":"Department of Agricultural Biotechnology Seoul National University Seoul South Korea"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2023,7,1]]},"reference":[{"key":"2024012019384007700_jeb14153-bib-0001","doi-asserted-by":"crossref","first-page":"762","DOI":"10.1086\/668011","article-title":"A quantitative test of the predicted relationship between countershading and lighting environment","volume":"180","author":"Allen","year":"2012","journal-title":"The American Naturalist"},{"key":"2024012019384007700_jeb14153-bib-0002","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1080\/03949370.2016.1145146","article-title":"Adaptive significance of coat colouration and patterns of Sciuromorpha (Rodentia)","volume":"29","author":"Ancillotto","year":"2017","journal-title":"Ethology Ecology and Evolution"},{"key":"2024012019384007700_jeb14153-bib-0003","doi-asserted-by":"crossref","first-page":"13597","DOI":"10.1073\/pnas.1509811112","article-title":"Antipredator defenses predict diversification rates","volume":"112","author":"Arbuckle","year":"2015","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2024012019384007700_jeb14153-bib-0004","doi-asserted-by":"crossref","first-page":"2812","DOI":"10.1073\/pnas.1421926112","article-title":"Moth tails divert bat attack: Evolution of acoustic deflection","volume":"112","author":"Barber","year":"2015","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2024012019384007700_jeb14153-bib-0005","doi-asserted-by":"crossref","first-page":"590","DOI":"10.2307\/1369048","article-title":"Melanin and the abrasion resistance of feathers","volume":"97","author":"Bonser","year":"1995","journal-title":"Condor"},{"key":"2024012019384007700_jeb14153-bib-0006","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1017\/S0952836901000322","article-title":"The ontogeny and distribution of countershading in colonies of the naked mole-rat (Heterocephalus glaber )","volume":"253","author":"Braude","year":"2001","journal-title":"Journal of Zoology"},{"key":"2024012019384007700_jeb14153-bib-0007","doi-asserted-by":"crossref","first-page":"1231","DOI":"10.1007\/s10682-011-9479-5","article-title":"The functional significance of colouration in cetaceans","volume":"25","author":"Caro","year":"2011","journal-title":"Evolutionary Ecology"},{"key":"2024012019384007700_jeb14153-bib-0008","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1111\/brv.12670","article-title":"Towards an ecology of protective coloration","volume":"96","author":"Caro","year":"2021","journal-title":"Biological Reviews"},{"key":"2024012019384007700_jeb14153-bib-0009","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1093\/beheco\/ars025","article-title":"Pelage coloration in pinnipeds: Functional considerations","volume":"23","author":"Caro","year":"2012","journal-title":"Behavioral Ecology"},{"volume-title":"Adaptive coloration in animals","year":"1940","author":"Cott","key":"2024012019384007700_jeb14153-bib-0010"},{"key":"2024012019384007700_jeb14153-bib-0011","doi-asserted-by":"crossref","first-page":"13093","DOI":"10.1073\/pnas.1611589113","article-title":"Optimizing countershading camouflage","volume":"113","author":"Cuthill","year":"2016","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"2024012019384007700_jeb14153-bib-0012","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1163\/036551656X00012","article-title":"Countershading in caterpillars","volume":"11","author":"De Ruiter","year":"1956","journal-title":"Archives N\u00e9erlandaises de Zoologie"},{"key":"2024012019384007700_jeb14153-bib-0013","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2020.0477","volume-title":"Countershading enhances camouflage by reducing prey contrast: Countershading reduces detection","author":"Donohue","year":"2020"},{"key":"2024012019384007700_jeb14153-bib-0014","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1111\/j.1095-8312.1994.tb00973.x","article-title":"The survival value of countershading with wild birds as predators","volume":"51","author":"Edmunds","year":"1994","journal-title":"Biological Journal of the Linnean Society"},{"key":"2024012019384007700_jeb14153-bib-0015","first-page":"10","article-title":"Ultraviolet photography of forty-three species of lepidoptera representing ten families","volume":"56","author":"Eguchi","year":"1983","journal-title":"Annotationes Zoologicae Japonenses"},{"key":"2024012019384007700_jeb14153-bib-0016","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1007\/978-1-4615-6956-5_5","volume-title":"Evolutionary biology","author":"Endler","year":"1978"},{"key":"2024012019384007700_jeb14153-bib-0017","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1098\/rspb.1991.0011","article-title":"A countershading reflex in cephalopods","volume":"243","author":"Ferguson","year":"1991","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"key":"2024012019384007700_jeb14153-bib-0018","doi-asserted-by":"crossref","first-page":"S42","DOI":"10.1086\/510138","article-title":"Simultaneous crypsis and conspicuousness in color patterns: Comparative analysis of a neotropical rainforest bird community","volume":"169","author":"Gomez","year":"2007","journal-title":"The American Naturalist"},{"key":"2024012019384007700_jeb14153-bib-0019","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/S0003-3472(71)80017-9","article-title":"Countershading in the colourful reef fish Chaetodon lunula: Concealment, communication or both?","volume":"19","author":"Hamilton","year":"1971","journal-title":"Animal Behaviour"},{"key":"2024012019384007700_jeb14153-bib-0021","doi-asserted-by":"crossref","first-page":"6664","DOI":"10.1073\/pnas.1415121112","article-title":"Body size affects the evolution of eyespots in caterpillars","volume":"112","author":"Hossie","year":"2015","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2024012019384007700_jeb14153-bib-0022","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1007\/s10764-009-9359-9","article-title":"Interspecific variation in primate countershading: Effects of activity pattern, body mass, and phylogeny","volume":"30","author":"Kamilar","year":"2009","journal-title":"International Journal of Primatology"},{"key":"2024012019384007700_jeb14153-bib-0023","doi-asserted-by":"crossref","first-page":"20171287","DOI":"10.1098\/rspb.2017.1287","article-title":"Body size affects the evolution of hidden colour signals in moths","volume":"284","author":"Kang","year":"2017","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"key":"2024012019384007700_jeb14153-bib-0024","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1111\/bij.12291","article-title":"Body size influences differently the detectabilities of colour morphs of cryptic prey","volume":"113","author":"Karpestam","year":"2014","journal-title":"Biological Journal of the Linnean Society"},{"key":"2024012019384007700_jeb14153-bib-0025","doi-asserted-by":"crossref","first-page":"e5719","DOI":"10.1371\/journal.pone.0005719","article-title":"Phylogeny and biogeography of hawkmoths (lepidoptera: Sphingidae): Evidence from five nuclear genes","volume":"4","author":"Kawahara","year":"2009","journal-title":"PLoS One"},{"key":"2024012019384007700_jeb14153-bib-0026","doi-asserted-by":"crossref","first-page":"20201894","DOI":"10.1098\/rspb.2020.1894","article-title":"Prey with hidden colour defences benefit from their similarity to aposematic signals","volume":"287","author":"Kim","year":"2020","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"key":"2024012019384007700_jeb14153-bib-0027","doi-asserted-by":"crossref","DOI":"10.1007\/b97377","volume-title":"Survival analysis: Techniques for censored and truncated data","author":"Klein","year":"2003"},{"key":"2024012019384007700_jeb14153-bib-0028","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1086\/703535","article-title":"Consistent associations between body size and hidden contrasting color signals across a range of insect taxa","volume":"194","author":"Loeffler-Henry","year":"2019","journal-title":"The American Naturalist"},{"key":"2024012019384007700_jeb14153-bib-0029","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1007\/s10682-006-9130-z","article-title":"Size dependent predation risk in cryptic and conspicuous insects","volume":"21","author":"M\u00e4nd","year":"2007","journal-title":"Evolutionary Ecology"},{"key":"2024012019384007700_jeb14153-bib-0030","doi-asserted-by":"crossref","first-page":"5016","DOI":"10.1038\/ncomms6016","article-title":"Seasonal changes in predator community switch the direction of selection for prey defences","volume":"5","author":"Mappes","year":"2014","journal-title":"Nature Communications"},{"key":"2024012019384007700_jeb14153-bib-0031","doi-asserted-by":"crossref","first-page":"e7386","DOI":"10.7717\/peerj.7386","article-title":"A comprehensive molecular phylogeny of Geometridae (lepidoptera) with a focus on enigmatic small subfamilies","volume":"2019","author":"Murillo-Ramos","year":"2019","journal-title":"PeerJ"},{"key":"2024012019384007700_jeb14153-bib-0032","doi-asserted-by":"crossref","first-page":"2042","DOI":"10.1016\/j.visres.2008.06.018","article-title":"A review of the evolution of animal colour vision and visual communication signals","volume":"48","author":"Osorio","year":"2008","journal-title":"Vision Research"},{"key":"2024012019384007700_jeb14153-bib-0033","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1093\/bioinformatics\/btg412","article-title":"APE: Analyses of phylogenetics and evolution in R language","volume":"20","author":"Paradis","year":"2004","journal-title":"Bioinformatics"},{"key":"2024012019384007700_jeb14153-bib-0034","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1007\/s10164-021-00691-5","article-title":"Size-dependent predation risk in cryptic prey","volume":"39","author":"Pembury Smith","year":"2021","journal-title":"Journal of Ethology"},{"key":"2024012019384007700_jeb14153-bib-0035","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1111\/1365-2435.12481","article-title":"Orientation to the sun by animals and its interaction with crypsis","volume":"29","author":"Penacchio","year":"2015","journal-title":"Functional Ecology"},{"key":"2024012019384007700_jeb14153-bib-0036","doi-asserted-by":"crossref","first-page":"170801","DOI":"10.1098\/rsos.170801","article-title":"Is countershading camouflage robust to lighting change due to weather?","volume":"5","author":"Penacchio","year":"2018","journal-title":"Royal Society Open Science"},{"key":"2024012019384007700_jeb14153-bib-0037","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1038\/nature10961","article-title":"A comparative analysis of the evolution of imperfect mimicry","volume":"483","author":"Penney","year":"2012","journal-title":"Nature"},{"volume-title":"The colours of animals: Their meaning and use, especially considered in the case of insects","year":"1890","author":"Poulton","key":"2024012019384007700_jeb14153-bib-0038"},{"key":"2024012019384007700_jeb14153-bib-0039","doi-asserted-by":"crossref","first-page":"973","DOI":"10.1111\/j.1365-2656.2009.01566.x","article-title":"Size-dependent predation risk in tree-feeding insects with different colouration strategies: A field experiment","volume":"78","author":"Remmel","year":"2009","journal-title":"The Journal of Animal Ecology"},{"key":"2024012019384007700_jeb14153-bib-0040","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1098\/rstb.2008.0261","article-title":"From Abbott Thayer to the present day: What have we learned about the function of countershading?","volume":"364","author":"Rowland","year":"2009","journal-title":"Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences"},{"key":"2024012019384007700_jeb14153-bib-0041","doi-asserted-by":"crossref","first-page":"2539","DOI":"10.1098\/rspb.2008.0812","article-title":"Can't tell the caterpillars from the trees: Countershading enhances survival in a woodland","volume":"275","author":"Rowland","year":"2008","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"key":"2024012019384007700_jeb14153-bib-0042","doi-asserted-by":"crossref","first-page":"1249","DOI":"10.1016\/j.anbehav.2007.01.030","article-title":"Countershading enhances cryptic protection: An experiment with wild birds and artificial prey","volume":"74","author":"Rowland","year":"2007","journal-title":"Animal Behaviour"},{"key":"2024012019384007700_jeb14153-bib-0043","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780199688678.001.0001","volume-title":"Avoiding attack: The evolutionary ecology of crypsis, aposematism, and mimicry","author":"Ruxton","year":"2018","edition":"2nd ed."},{"key":"2024012019384007700_jeb14153-bib-0044","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1016\/j.anbehav.2003.12.009","article-title":"What, if anything, is the adaptive function of countershading?","volume":"68","author":"Ruxton","year":"2004","journal-title":"Animal Behaviour"},{"key":"2024012019384007700_jeb14153-bib-0045","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1093\/oxfordjournals.molbev.a003974","article-title":"Estimating absolute rates of molecular evolution and divergence times: A penalized likelihood approach","volume":"19","author":"Sanderson","year":"2002","journal-title":"Molecular Biology and Evolution"},{"key":"2024012019384007700_jeb14153-bib-0046","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1126\/science.1181931","article-title":"Masquerade: Camouflage without crypsis","volume":"327","author":"Skelhorn","year":"2010","journal-title":"Science"},{"key":"2024012019384007700_jeb14153-bib-0047","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1111\/brv.12438","article-title":"The key role of behaviour in animal camouflage","volume":"94","author":"Stevens","year":"2019","journal-title":"Biological Reviews"},{"key":"2024012019384007700_jeb14153-bib-0048","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1093\/beheco\/arg072","article-title":"Ecological and behavioral correlates of coloration in artiodactyls: Systematic analyses of conventional hypotheses","volume":"14","author":"Stoner","year":"2003","journal-title":"Behavioral Ecology"},{"key":"2024012019384007700_jeb14153-bib-0049","doi-asserted-by":"crossref","first-page":"124","DOI":"10.2307\/4068693","article-title":"The law which underlies protective coloration","volume":"13","author":"Thayer","year":"1896","journal-title":"Auk"},{"volume-title":"Concealing-coloration in the animal kingdom: An exposition of the laws of disguise through color and pattern","year":"1918","author":"Thayer","key":"2024012019384007700_jeb14153-bib-0050"},{"first-page":"1","year":"2020","author":"Therneau","key":"2024012019384007700_jeb14153-bib-0051"},{"key":"2024012019384007700_jeb14153-bib-0020","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1093\/sysbio\/syu005","article-title":"A linear-time algorithm for gaussian and non-gaussian trait evolution models","volume":"63","author":"Tung Ho","year":"2014","journal-title":"Systematic Biology"},{"key":"2024012019384007700_jeb14153-bib-0052","doi-asserted-by":"crossref","first-page":"1814","DOI":"10.1016\/S0003-3472(87)80074-X","article-title":"Thermoregulation and daily activity patterns in a black desert grasshopper, Taeniopoda eques","volume":"35","author":"Whitman","year":"1987","journal-title":"Animal Behaviour"}],"container-title":["Journal of Evolutionary Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/jeb.14153","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/jeb.14153","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/academic.oup.com\/jeb\/article-pdf\/36\/7\/1032\/56276028\/jeb14153.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jeb\/article-pdf\/36\/7\/1032\/56276028\/jeb14153.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,1,23]],"date-time":"2024-01-23T20:33:17Z","timestamp":1706041997000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jeb\/article\/36\/7\/1032-1039\/7577255"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,7,1]]},"references-count":52,"journal-issue":{"issue":"7","published-print":{"date-parts":[[2023,7,1]]}},"URL":"https:\/\/doi.org\/10.1111\/jeb.14153","relation":{},"ISSN":["1420-9101","1010-061X"],"issn-type":[{"type":"electronic","value":"1420-9101"},{"type":"print","value":"1010-061X"}],"subject":[],"published-other":{"date-parts":[[2023,7,1]]},"published":{"date-parts":[[2023,7,1]]},"assertion":[{"value":"2022-05-09","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-12-14","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-02-03","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}