{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,4]],"date-time":"2026-07-04T23:16:17Z","timestamp":1783206977353,"version":"3.54.6"},"reference-count":49,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T00:00:00Z","timestamp":1781136000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/100014809","name":"Technology Agency of the Czech Republic","doi-asserted-by":"publisher","award":["SS07010197"],"award-info":[{"award-number":["SS07010197"]}],"id":[{"id":"10.13039\/100014809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000921","name":"European Cooperation in Science and Technology","doi-asserted-by":"publisher","award":["014\/2022\/P"],"award-info":[{"award-number":["014\/2022\/P"]}],"id":[{"id":"10.13039\/501100000921","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000921","name":"European Cooperation in Science and Technology","doi-asserted-by":"publisher","award":["LUC25115"],"award-info":[{"award-number":["LUC25115"]}],"id":[{"id":"10.13039\/501100000921","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001823","name":"Ministerstvo \u0160kolstv\u00ed, Ml\u00e1de\u017ee a T\u011blov\u00fdchovy","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001823","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001824","name":"Czech Science Foundation","doi-asserted-by":"publisher","award":["26-22878S"],"award-info":[{"award-number":["26-22878S"]}],"id":[{"id":"10.13039\/501100001824","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Journal of Thermal Biology"],"published-print":{"date-parts":[[2026,7]]},"DOI":"10.1016\/j.jtherbio.2026.104497","type":"journal-article","created":{"date-parts":[[2026,5,29]],"date-time":"2026-05-29T15:54:16Z","timestamp":1780070056000},"page":"104497","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["Heated debate: Is the seasonal polyphenism of the map butterfly Araschnia levana driven by thermoregulation?"],"prefix":"10.1016","volume":"139","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0686-3961","authenticated-orcid":false,"given":"Daniel","family":"Linke","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9935-7859","authenticated-orcid":false,"given":"Jan","family":"Okrouhlik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6298-2466","authenticated-orcid":false,"given":"Alena Such\u00e1\u010dkov\u00e1","family":"Barto\u0148ov\u00e1","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4518-4328","authenticated-orcid":false,"given":"Leonardo","family":"R\u00e9 Jorge","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2885-4919","authenticated-orcid":false,"given":"P\u00e1vel","family":"Matos-Marav\u00ed","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9333-213X","authenticated-orcid":false,"given":"Irena","family":"Kleckova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jtherbio.2026.104497_bib1","doi-asserted-by":"crossref","DOI":"10.7717\/peerj.3349","article-title":"Exoskeleton may influence the internal body temperatures of neotropical dung beetles (Col. Scarabaeinae)","volume":"5","author":"Amore","year":"2017","journal-title":"PeerJ"},{"issue":"9","key":"10.1016\/j.jtherbio.2026.104497_bib2","doi-asserted-by":"crossref","first-page":"1759","DOI":"10.1111\/1365-2656.13970","article-title":"Tropical butterflies use thermal buffering and thermal tolerance as alternative strategies to cope with temperature increase","volume":"92","author":"Ashe-Jepson","year":"2023","journal-title":"J. Anim. Ecol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v067.i01","article-title":"Fitting linear mixed-effects models using lme4","volume":"67","author":"Bates","year":"2015","journal-title":"J. Stat. Software"},{"issue":"10","key":"10.1016\/j.jtherbio.2026.104497_bib4","doi-asserted-by":"crossref","first-page":"4891","DOI":"10.1002\/ece3.4047","article-title":"Immunological larval polyphenism in the map butterfly Araschnia levana reveals the photoperiodic modulation of immunity","volume":"8","author":"Baudach","year":"2018","journal-title":"Ecol. Evol."},{"issue":"11","key":"10.1016\/j.jtherbio.2026.104497_bib5","doi-asserted-by":"crossref","first-page":"2440","DOI":"10.1111\/1365-2656.13319","article-title":"How butterflies keep their cool: physical and ecological traits influence thermoregulatory ability and population trends","volume":"89","author":"Bladon","year":"2020","journal-title":"J. Anim. Ecol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib6","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.jtherbio.2016.01.007","article-title":"Cool bands: wing bands decrease rate of heating, but not equilibrium temperature in Anartia fatima","volume":"56","author":"Brashears","year":"2016","journal-title":"J. Therm. Biol."},{"issue":"368","key":"10.1016\/j.jtherbio.2026.104497_bib7","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1080\/01621459.1979.10481038","article-title":"Robust locally weighted regression and smoothing scatterplots","volume":"74","author":"Cleveland","year":"1979","journal-title":"J. Am. Stat. Assoc."},{"issue":"2","key":"10.1016\/j.jtherbio.2026.104497_bib8","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1111\/j.1365-2435.2007.01377.x","article-title":"Testing the thermal melanism hypothesis: a macrophysiological approach","volume":"22","author":"Clusella\u2010Trullas","year":"2008","journal-title":"Funct. Ecol."},{"issue":"1","key":"10.1016\/j.jtherbio.2026.104497_bib9","doi-asserted-by":"crossref","first-page":"1294","DOI":"10.1038\/s41467-020-15033-1","article-title":"Diverse nanostructures underlie thin ultra-black scales in butterflies","volume":"11","author":"Davis","year":"2020","journal-title":"Nat. Commun."},{"issue":"4","key":"10.1016\/j.jtherbio.2026.104497_bib10","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0122623","article-title":"Why small is beautiful: wing colour is free from thermoregulatory constraint in the small lycaenid butterfly, polyommatus icarus","volume":"10","author":"De Keyser","year":"2015","journal-title":"PLoS One"},{"issue":"6","key":"10.1016\/j.jtherbio.2026.104497_bib11","first-page":"1601","article-title":"Seasonal polyphenism in Bicyclus dorothea (Lepidoptera: nymphalidae) across different habitats in Cameroon","volume":"47","author":"Dongmo","year":"2018","journal-title":"Environ. Entomol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib12","doi-asserted-by":"crossref","DOI":"10.3389\/fevo.2021.612330","article-title":"Ontogenetic basis of among-generation differences in size-related traits in a polyphenic butterfly","volume":"9","author":"Esperk","year":"2021","journal-title":"Front. Ecol. Evol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib13","first-page":"603","article-title":"Life-history polyphenism in the map butterfly (Araschnia levana): developmental constraints versus season-specific adaptations","volume":"12","author":"Friberg","year":"2010","journal-title":"Evol. Ecol. Res."},{"issue":"8","key":"10.1016\/j.jtherbio.2026.104497_bib14","first-page":"1511","article-title":"Mechanical design indicates differences in mobility among butterfly generations","volume":"8","author":"Fric","year":"2006","journal-title":"Evol. Ecol. Res."},{"issue":"2","key":"10.1016\/j.jtherbio.2026.104497_bib15","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1111\/j.1420-9101.2003.00681.x","article-title":"Red & black or black & white? Phylogeny of the Araschnia butterflies (Lepidoptera: nymphalidae) and evolution of seasonal polyphenism","volume":"17","author":"Fric","year":"2004","journal-title":"J. Evol. Biol."},{"issue":"7","key":"10.1016\/j.jtherbio.2026.104497_bib16","first-page":"1017","article-title":"Generations of the polyphenic butterfly Araschnia levana differ in body design","volume":"4","author":"Fric","year":"2002","journal-title":"Evol. Ecol. Res."},{"issue":"7","key":"10.1016\/j.jtherbio.2026.104497_bib17","doi-asserted-by":"crossref","first-page":"1302","DOI":"10.1093\/evolut\/qpae059","article-title":"Phenotypic plasticity in tropical butterflies is linked to climatic seasonality on a macroevolutionary scale","volume":"78","author":"Halali","year":"2024","journal-title":"Evolution"},{"issue":"1","key":"10.1016\/j.jtherbio.2026.104497_bib18","first-page":"34","article-title":"Seasonal plasticity in Junonia coenia (Nymphalidae): linking wing color, temperature dynamics, and behavior","volume":"73","author":"J\u00e4rvi","year":"2019","journal-title":"J. Lepid. Soc."},{"issue":"6","key":"10.1016\/j.jtherbio.2026.104497_bib19","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1046\/j.1420-9101.2002.00470.x","article-title":"A novel method of behavioural thermoregulation in butterflies","volume":"15","author":"Kemp","year":"2002","journal-title":"J. Evol. Biol."},{"issue":"5","key":"10.1016\/j.jtherbio.2026.104497_bib20","doi-asserted-by":"crossref","first-page":"942","DOI":"10.1111\/j.1558-5646.1995.tb02329.x","article-title":"Fitness consequences of seasonal polyphenism in Western white butterflies","volume":"49","author":"Kingsolver","year":"1995","journal-title":"Evolution"},{"key":"10.1016\/j.jtherbio.2026.104497_bib21","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.jtherbio.2014.02.002","article-title":"Thermoregulation and microhabitat use in mountain butterflies of the genus Erebia: importance of fine-scale habitat heterogeneity","volume":"41","author":"Kleckova","year":"2014","journal-title":"J. Therm. Biol."},{"issue":"7","key":"10.1016\/j.jtherbio.2026.104497_bib22","doi-asserted-by":"crossref","DOI":"10.1002\/ece3.70012","article-title":"Flight behaviour diverges more between seasonal forms than between species in Pieris butterflies","volume":"14","author":"Kleckova","year":"2024","journal-title":"Ecol. Evol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib23","doi-asserted-by":"crossref","DOI":"10.1016\/j.jtherbio.2023.103502","article-title":"Body size, not species identity, drives body heating in alpine Erebia butterflies","volume":"113","author":"Kleckova","year":"2023","journal-title":"J. Therm. Biol."},{"issue":"1","key":"10.1016\/j.jtherbio.2026.104497_bib24","first-page":"365","article-title":"Seasonal polyphenism in body size and juvenile development of the swallowtail butterfly Papilio xuthus (Lepidoptera: papilionidae)","volume":"114","author":"Komata","year":"2017","journal-title":"EJE (Eur. J. Epidemiol.)"},{"issue":"18","key":"10.1016\/j.jtherbio.2026.104497_bib25","doi-asserted-by":"crossref","DOI":"10.1242\/jeb.245751","article-title":"Recent advances in insect thermoregulation","volume":"226","author":"Lahond\u00e8re","year":"2023","journal-title":"J. Exp. Biol."},{"issue":"15","key":"10.1016\/j.jtherbio.2026.104497_bib26","doi-asserted-by":"crossref","first-page":"4180","DOI":"10.1111\/gcb.16797","article-title":"Thermoregulatory ability and mechanism do not differ consistently between neotropical and temperate butterflies","volume":"29","author":"Laird-Hopkins","year":"2023","journal-title":"Glob. Change Biol."},{"issue":"9","key":"10.1016\/j.jtherbio.2026.104497_bib27","doi-asserted-by":"crossref","first-page":"1054","DOI":"10.1039\/D2NH00166G","article-title":"Biomimetic reconstruction of butterfly wing scale nanostructures for radiative cooling and structural coloration","volume":"7","author":"Lee","year":"2022","journal-title":"Nanoscale Horiz."},{"key":"10.1016\/j.jtherbio.2026.104497_bib28","author":"Lenth"},{"key":"10.1016\/j.jtherbio.2026.104497_bib29","series-title":"Association of Iridescence and Tails with Flight Proxies Suggests that Predation Drives Convergence in Skipper Butterflies","first-page":"2025","author":"Linke","year":"2025"},{"issue":"6","key":"10.1016\/j.jtherbio.2026.104497_bib30","doi-asserted-by":"crossref","DOI":"10.1002\/ece3.8992","article-title":"Thermobiological effects of temperature-induced color variations in Aglais urticae (Lepidoptera, nymphalidae)","volume":"12","author":"Markl","year":"2022","journal-title":"Ecol. Evol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib31","series-title":"Thermal Biology: Temperature Sensing and Temperature-Responding Systems","first-page":"79","article-title":"Lipid-mediated mechanisms of thermal adaptation and thermoregulatory behavior in animals","author":"Nagao","year":"2024"},{"key":"10.1016\/j.jtherbio.2026.104497_bib32","series-title":"Photoperiodism: the Biological Calendar","author":"Nelson","year":"2010"},{"key":"10.1016\/j.jtherbio.2026.104497_bib33","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1016\/j.jtherbio.2018.07.014","article-title":"Photonic structures improve radiative heat exchange of Rosalia alpina (Coleoptera: cerambycidae)","volume":"76","author":"Pavlovic","year":"2018","journal-title":"J. Therm. Biol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib34","series-title":"Nlme: Linear and Nonlinear Mixed Effects Models","author":"Pinheiro","year":"2025"},{"key":"10.1016\/j.jtherbio.2026.104497_bib35","series-title":"R: a Language and Environment for Statistical Computing","year":"2025"},{"key":"10.1016\/j.jtherbio.2026.104497_bib36","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.cbpb.2017.11.017","article-title":"Diversity in membrane composition is associated with variation in thermoregulatory capacity in hymenopterans","volume":"224","author":"Rodr\u00edguez","year":"2018","journal-title":"Comp. Biochem. Physiol. B Biochem. Mol. Biol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib37","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1016\/j.anbehav.2020.07.015","article-title":"Beyond thermal melanism: association of wing melanization with fitness and flight behaviour in a butterfly","volume":"167","author":"Rosa","year":"2020","journal-title":"Anim. Behav."},{"issue":"5","key":"10.1016\/j.jtherbio.2026.104497_bib38","doi-asserted-by":"crossref","first-page":"1208","DOI":"10.1093\/icb\/icaa124","article-title":"Circulation in insect wings","volume":"60","author":"Salcedo","year":"2020","journal-title":"Integr. Comp. Biol."},{"issue":"7","key":"10.1016\/j.jtherbio.2026.104497_bib39","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1038\/nmeth.2089","article-title":"NIH image to ImageJ: 25 years of image analysis","volume":"9","author":"Schneider","year":"2012","journal-title":"Nat. Methods"},{"issue":"1","key":"10.1016\/j.jtherbio.2026.104497_bib40","doi-asserted-by":"crossref","first-page":"174","DOI":"10.1111\/j.1365-2656.2011.01909.x","article-title":"A meta-analysis of the traits affecting dispersal ability in butterflies: can wingspan be used as a proxy?","volume":"81","author":"Sekar","year":"2012","journal-title":"J. Anim. Ecol."},{"issue":"5","key":"10.1016\/j.jtherbio.2026.104497_bib41","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1111\/1365-2435.12568","article-title":"Tropical to subpolar gradient in phospholipid composition suggests adaptive tuning of biological membrane function in drosophilids","volume":"30","author":"Slotsbo","year":"2016","journal-title":"Funct. Ecol."},{"issue":"14","key":"10.1016\/j.jtherbio.2026.104497_bib42","doi-asserted-by":"crossref","first-page":"7669","DOI":"10.1002\/ece3.6491","article-title":"Seasonal polyphenism of spotted\u2010wing drosophila is affected by variation in local abiotic conditions within its invaded range, likely influencing survival and regional population dynamics","volume":"10","author":"Stockton","year":"2020","journal-title":"Ecol. Evol."},{"issue":"6","key":"10.1016\/j.jtherbio.2026.104497_bib43","doi-asserted-by":"crossref","first-page":"1761","DOI":"10.1111\/1744-7917.13046","article-title":"Universal cooling patterns of the butterfly wing scales hierarchy deduced from the heterogeneous thermal and structural properties of Tirumala limniace (Lepidoptera: nymphalidae, Danainae)","volume":"29","author":"Tang","year":"2022","journal-title":"Insect Sci."},{"issue":"1","key":"10.1016\/j.jtherbio.2026.104497_bib44","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1038\/s41467-020-14408-8","article-title":"Physical and behavioral adaptations to prevent overheating of the living wings of butterflies","volume":"11","author":"Tsai","year":"2020","journal-title":"Nat. Commun."},{"issue":"2","key":"10.1016\/j.jtherbio.2026.104497_bib45","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1046\/j.1420-9101.2002.00384.x","article-title":"Seasonal butterfly design: morphological plasticity among three developmental pathways relative to sex, flight and thermoregulation","volume":"15","author":"Van Dyck","year":"2002","journal-title":"J. Evol. Biol."},{"issue":"19","key":"10.1016\/j.jtherbio.2026.104497_bib46","doi-asserted-by":"crossref","DOI":"10.1242\/jeb.242898","article-title":"Heating rates are more strongly influenced by near-infrared than visible reflectance in beetles","volume":"224","author":"Wang","year":"2021","journal-title":"J. Exp. Biol."},{"issue":"1","key":"10.1016\/j.jtherbio.2026.104497_bib47","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1007\/BF00689287","article-title":"Antagonism between haemolymph transport and tracheal ventilation in an insect wing (Attacus atlas L.)","volume":"147","author":"Wasserthal","year":"1982","journal-title":"J. Comp. Physiol."},{"key":"10.1016\/j.jtherbio.2026.104497_bib48","series-title":"\"Programming with ggplot2.\" Ggplot2: elegant graphics for data analysis","first-page":"241","author":"Wickham","year":"2016"},{"issue":"22","key":"10.1016\/j.jtherbio.2026.104497_bib49","doi-asserted-by":"crossref","first-page":"8062","DOI":"10.1002\/ece3.2464","article-title":"Cool habitats support darker and bigger butterflies in Australian tropical forests","volume":"6","author":"Xing","year":"2016","journal-title":"Ecol. Evol."}],"container-title":["Journal of Thermal Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0306456526001300?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0306456526001300?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,7,4]],"date-time":"2026-07-04T22:27:31Z","timestamp":1783204051000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0306456526001300"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7]]},"references-count":49,"alternative-id":["S0306456526001300"],"URL":"https:\/\/doi.org\/10.1016\/j.jtherbio.2026.104497","relation":{},"ISSN":["0306-4565"],"issn-type":[{"value":"0306-4565","type":"print"}],"subject":[],"published":{"date-parts":[[2026,7]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Heated debate: Is the seasonal polyphenism of the map butterfly Araschnia levana driven by thermoregulation?","name":"articletitle","label":"Article Title"},{"value":"Journal of Thermal Biology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jtherbio.2026.104497","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier Ltd.","name":"copyright","label":"Copyright"}],"article-number":"104497"}}