{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T17:42:00Z","timestamp":1789753320735,"version":"4.0.1"},"reference-count":123,"publisher":"Oxford University Press (OUP)","issue":"6","license":[{"start":{"date-parts":[[2026,5,30]],"date-time":"2026-05-30T00:00:00Z","timestamp":1780099200000},"content-version":"vor","delay-in-days":1,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["1145978"],"award-info":[{"award-number":["1145978"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Gopher Tortoise Council"},{"DOI":"10.13039\/100006597","name":"Florida State University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006597","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008900","name":"University of South Florida","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100008900","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,6,2]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Rapid adaptive evolution may be more likely to occur not only through standing genetic variation but via existing axes of genetic variation that have previously been exposed to selection. Ontogenetic variation represents one such axis and often evolves under strong selection in snake venoms. Snake venoms are complex cocktails of proteinaceous toxins, and ontogenetic shifts in venom expression are frequent and reflect dietary shifts across life history. Here, we used morphological, proteomic, transcriptomic, epigenomic, and optical genome mapping data to investigate a well-studied island-mainland population pair of eastern diamondback rattlesnakes (Crotalus adamanteus) to determine whether rapid adaptive expression divergence across populations occurred through the co-option of the ontogenetic regulatory network, population-specific changes independent of ontogeny, or a combination thereof. We found that island snakes were significantly smaller than mainland individuals, and venom proteomic data showed that the continuous ontogenetic shift in venom expression in the mainland population was truncated in island snakes. Venom-gland RNA-seq showed that island adults exhibited juvenile-like expression patterns at key transcription factors, and chromatin accessibility was predictive of venom gene differential expression for ontogenetically co-opted venom loci. Overall, rapid adaptation in the island population appears to have predominantly occurred through the co-option and truncation of the ontogenetic venom shift, with spatial differentiation playing a secondary role. Comparative tests in other systems are needed to determine whether rapid adaptation in general is not only biased toward standing genetic variation but toward large pre-existing axes of variation that have or continue to evolve under strong selection.<\/jats:p>","DOI":"10.1093\/gbe\/evag131","type":"journal-article","created":{"date-parts":[[2026,5,29]],"date-time":"2026-05-29T11:45:00Z","timestamp":1780055100000},"source":"Crossref","is-referenced-by-count":1,"title":["Truncated Life History Underlies Rapid Local Adaptation in Island Rattlesnake Venom Expression"],"prefix":"10.1093","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6153-6701","authenticated-orcid":false,"given":"Mark J","family":"Margres","sequence":"first","affiliation":[{"name":"Department of Integrative Biology, University of South Florida , Tampa, FL 33620 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3172-5210","authenticated-orcid":false,"given":"Samuel R","family":"Hirst","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of South Florida , Tampa, FL 33620 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-5246-0666","authenticated-orcid":false,"given":"Dylan G","family":"Gallinson","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of South Florida , Tampa, FL 33620 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8229-5951","authenticated-orcid":false,"given":"Rhett M","family":"Rautsaw","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of South Florida , Tampa, FL 33620 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7153-4611","authenticated-orcid":false,"given":"Preston J","family":"McDonald","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of South Florida , Tampa, FL 33620 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2601-1253","authenticated-orcid":false,"given":"Ella G","family":"Guedouar","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of South Florida , Tampa, FL 33620 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5388-3634","authenticated-orcid":false,"given":"Gunnar S","family":"Nystrom","sequence":"additional","affiliation":[{"name":"Department of Biological Science, Florida State University , Tallahassee, FL 32306 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3702-5079","authenticated-orcid":false,"given":"Michael P","family":"Hogan","sequence":"additional","affiliation":[{"name":"Department of Biological Science, Florida State University , Tallahassee, FL 32306 ,","place":["USA"]},{"name":"Department of Ecology & Evolutionary Biology, University of Michigan , Ann Arbor, MI 48109 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8321-1158","authenticated-orcid":false,"given":"Schyler A","family":"Ellsworth","sequence":"additional","affiliation":[{"name":"Department of Biological Science, Florida State University , Tallahassee, FL 32306 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6151-7703","authenticated-orcid":false,"given":"Kenneth P","family":"Wray","sequence":"additional","affiliation":[{"name":"Biodiversity Center, University of Texas at Austin , Austin, TX 78703 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0356-2178","authenticated-orcid":false,"given":"Darin R","family":"Rokyta","sequence":"additional","affiliation":[{"name":"Department of Biological Science, Florida State University , Tallahassee, FL 32306 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,5,29]]},"reference":[{"key":"2026060909570848800_evag131-B1","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","article-title":"HTSeq\u2014a Python framework to work with high-throughput sequencing data","volume":"31","author":"Anders","year":"2015","journal-title":"Bioinformatics"},{"key":"2026060909570848800_evag131-B2","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1126\/science.1159978","article-title":"Natural selection on a major armor gene in threespine stickleback","volume":"322","author":"Barrett","year":"2008","journal-title":"Science"},{"key":"2026060909570848800_evag131-B3","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1016\/j.tree.2007.09.008","article-title":"Adaptation from standing genetic variation","volume":"23","author":"Barrett","year":"2007","journal-title":"Trends Ecol Evol"},{"key":"2026060909570848800_evag131-B4","author":"Beerli"},{"key":"2026060909570848800_evag131-B5","author":"Blighe"},{"key":"2026060909570848800_evag131-B6","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1111\/j.0014-3820.2003.tb00268.x","article-title":"Empirical evidence for an optimal body size in snakes","volume":"57","author":"Boback","year":"2003","journal-title":"Evolution"},{"key":"2026060909570848800_evag131-B7","doi-asserted-by":"publisher","first-page":"110129","DOI":"10.1016\/j.cbpc.2025.110129","article-title":"Venom variation and ontogenetic changes in the Crotalus molossus complex: insights into composition, activities, and antivenom neutralization","volume":"290","author":"Borja","year":"2025","journal-title":"Comp Biochem Phys C"},{"key":"2026060909570848800_evag131-B8","doi-asserted-by":"publisher","first-page":"528","DOI":"10.1021\/pr9008749","article-title":"Snake venomics of the Central American rattlesnake Crotalus simus and the South American Crotalus durissus complex points to neurotoxicity as an adaptive paedomorphic trend along Crotalus dispersal in South America","volume":"9","author":"Calvete","year":"2010","journal-title":"J Prot Res"},{"key":"2026060909570848800_evag131-B9","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/0005-2795(78)90457-9","article-title":"Chemical and functional homology of myotoxin a from prairie rattlesnake venom and crotamine from South American rattlesnake venom","volume":"532","author":"Cameron","year":"1978","journal-title":"Biochim Biophys Acta"},{"key":"2026060909570848800_evag131-B10","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1111\/j.1365-2435.2007.01289.x","article-title":"Evolution on ecological time-scales","volume":"21","author":"Carroll","year":"2007","journal-title":"Funct Ecol"},{"key":"2026060909570848800_evag131-B11","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2307\/1936628","article-title":"A general explanation for insular body size trends in terrestrial vertebrates","volume":"59","author":"Case","year":"1978","journal-title":"Ecology"},{"key":"2026060909570848800_evag131-B12","doi-asserted-by":"publisher","first-page":"2637","DOI":"10.1093\/molbev\/msr091","article-title":"Domain loss facilitates accelerated evolution and neofunctionalization of duplicate snake venom metalloproteinase toxin genes","volume":"28","author":"Casewell","year":"2011","journal-title":"Mol Biol Evol"},{"key":"2026060909570848800_evag131-B13","doi-asserted-by":"publisher","first-page":"959","DOI":"10.1038\/nmeth.4396","article-title":"An improved ATAC-seq protocol reduces background and enables interrogation of frozen tissues","volume":"14","author":"Corces","year":"2017","journal-title":"Nat Methods"},{"key":"2026060909570848800_evag131-B14","doi-asserted-by":"publisher","first-page":"2156","DOI":"10.1093\/bioinformatics\/btr330","article-title":"The variant call format and VCFtools","volume":"27","author":"Danecek","year":"2011","journal-title":"Bioinformatics"},{"key":"2026060909570848800_evag131-B15","doi-asserted-by":"publisher","first-page":"giab008","DOI":"10.1093\/gigascience\/giab008","article-title":"Twelve years of samtools and bcftools","volume":"10","author":"Danecek","year":"2021","journal-title":"Gigascience"},{"key":"2026060909570848800_evag131-B16","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/ncomms9555","article-title":"Bayesian integration of genetics and epigenetics detects causal regulatory SNPs underlying expression variability","volume":"6","author":"Das","year":"2015","journal-title":"Nat Commun"},{"key":"2026060909570848800_evag131-B17","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1038\/nature08789","article-title":"Changes in Hox genes\u2019 structure and function during the evolution of the squamate body plan","volume":"464","author":"Di-Poi","year":"2010","journal-title":"Nature"},{"key":"2026060909570848800_evag131-B18","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1111\/evo.12284","article-title":"Why ontogeny matters during adaptation: developmental niche construction and pleiotropy across the life cycle in Arabidopsis thaliana","volume":"68","author":"Donohue","year":"2013","journal-title":"Evolution"},{"key":"2026060909570848800_evag131-B19","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/1471-2164-14-1","article-title":"Integrated \u201comics\u201d profiling indicates that miRNAs are modulators of the ontogenetic venom composition shift in the Central American rattlesnake, Crotalus simus simus","volume":"14","author":"Durban","year":"2013","journal-title":"BMC Genomics"},{"key":"2026060909570848800_evag131-B20","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1023\/A:1023818214614","article-title":"Isometric logratio transformations for compositional data analysis","volume":"35","author":"Egozcue","year":"2003","journal-title":"Math Geol"},{"key":"2026060909570848800_evag131-B21","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.23305","article-title":"ATAC-seq in emerging model organisms: challenges and strategies","author":"Erdo\u011fan","year":"2025","journal-title":"J Exp Zool Part B"},{"key":"2026060909570848800_evag131-B22","doi-asserted-by":"publisher","first-page":"e2409943121","DOI":"10.1073\/pnas.2409943121","article-title":"Fitness consequences of structural variation inferred from a house finch pangenome","volume":"121","author":"Fang","year":"2024","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026060909570848800_evag131-B23","doi-asserted-by":"publisher","first-page":"253","DOI":"10.2307\/5458","article-title":"Variation in sexual size dimorphism and maximum body size among adder populations: effects of prey size","volume":"60","author":"Forsman","year":"1991","journal-title":"J Anim Ecol"},{"key":"2026060909570848800_evag131-B24","doi-asserted-by":"publisher","first-page":"183","DOI":"10.2307\/2389885","article-title":"The advantage of a big head: swallowing performance in adders, Vipera berus","volume":"7","author":"Forsman","year":"1993","journal-title":"Funct Ecol"},{"key":"2026060909570848800_evag131-B25","doi-asserted-by":"publisher","first-page":"234","DOI":"10.1038\/202234a0","article-title":"Evolution of mammals on islands","volume":"202","author":"Foster","year":"1964","journal-title":"Nature"},{"key":"2026060909570848800_evag131-B26","doi-asserted-by":"publisher","first-page":"4065","DOI":"10.1242\/dev.129.17.4065","article-title":"Hoxa5 overexpression correlates with IGFBP1 upregulation and postnatal dwarfism: evidence for an interaction between Hoxa5 and Forkhead box transcription factors","volume":"129","author":"Foucher","year":"2002","journal-title":"Development"},{"key":"2026060909570848800_evag131-B27","doi-asserted-by":"publisher","first-page":"222","DOI":"10.2307\/1442480","article-title":"Reproduction, growth, and sexual dimorphism in the Canebrake Rattlesnake (Crotalus horridus atricaudatus)","volume":"1972","author":"Gibbons","year":"1972","journal-title":"Copeia"},{"key":"2026060909570848800_evag131-B28","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1007\/s00239-008-9186-1","article-title":"Snake population venomics: proteomics-based analyses of individual variation reveals significant gene regulation effects on venom protein expression in Sistrurus rattlesnakes","volume":"68","author":"Gibbs","year":"2009","journal-title":"J Mol Evol"},{"key":"2026060909570848800_evag131-B29","doi-asserted-by":"publisher","first-page":"evae110","DOI":"10.1093\/gbe\/evae110","article-title":"Diverse gene regulatory mechanisms alter rattlesnake venom gene expression at fine evolutionary scales","volume":"16","author":"Gopalan","year":"2024","journal-title":"Genome Biol Evol"},{"key":"2026060909570848800_evag131-B30","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1670\/22-068","article-title":"Heavy, bulky, or both: what does \u201clarge prey\u201d mean to snakes?","volume":"57","author":"Greene","year":"2023","journal-title":"J Herpetol"},{"key":"2026060909570848800_evag131-B31","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1038\/nature24277","article-title":"Genetic effects on gene expression across human tissues","volume":"550","author":"GTEx Consortium","year":"2017","journal-title":"Nature"},{"key":"2026060909570848800_evag131-B32","doi-asserted-by":"publisher","first-page":"109094","DOI":"10.1016\/j.toxicon.2026.109094","article-title":"Ontogenetic co-option of myotoxin expression variation in island eastern diamondback rattlesnake (crotalus adamanteus) venoms","volume":"277","author":"Guedouar","year":"2026","journal-title":"Toxicon"},{"key":"2026060909570848800_evag131-B33","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1016\/j.tig.2017.04.005","article-title":"Perspectives on gene regulatory network evolution","volume":"33","author":"Halfon","year":"2017","journal-title":"Trends Genet"},{"key":"2026060909570848800_evag131-B34","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/j.zool.2017.11.006","article-title":"Ontogenetic prey size selection in snakes: predator size and functional limitations to handling minimum prey sizes","volume":"126","author":"Hampton","year":"2018","journal-title":"Zoology"},{"key":"2026060909570848800_evag131-B35","doi-asserted-by":"publisher","first-page":"e1007859","DOI":"10.1371\/journal.pgen.1007859","article-title":"On the unfounded enthusiasm for soft selective sweeps ii: examining recent evidence from humans, flies, and viruses","volume":"14","author":"Harris","year":"2018","journal-title":"PLoS Genet"},{"key":"2026060909570848800_evag131-B36","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1038\/nature14042","article-title":"Evolution of the snake body form reveals homoplasy in amniote Hox gene function","volume":"520","author":"Head","year":"2015","journal-title":"Nature"},{"key":"2026060909570848800_evag131-B37","doi-asserted-by":"publisher","first-page":"evae198","DOI":"10.1093\/gbe\/evae198","article-title":"Where the \u2018ruber\u2019 meets the road: using the genome of the red diamond rattlesnake to unravel the evolutionary processes driving venom evolution","volume":"16","author":"Hirst","year":"2024","journal-title":"Genome Biol Evol"},{"key":"2026060909570848800_evag131-B38","doi-asserted-by":"publisher","first-page":"qpaf074","DOI":"10.1093\/evolut\/qpaf074","article-title":"Island biogeography and competition drive rapid venom complexity evolution across rattlesnakes","volume":"79","author":"Hirst","year":"2025","journal-title":"Evolution"},{"key":"2026060909570848800_evag131-B39","first-page":"76","article-title":"Lack of an ontogenetic shift in crotalus enyo cerralvensis venom supports integration of cranial morphology and venom expression in rattlesnakes","volume":"86","author":"Hirst","year":"2026","journal-title":"West N Am Nat"},{"key":"2026060909570848800_evag131-B40","doi-asserted-by":"publisher","first-page":"e2313440121","DOI":"10.1073\/pnas.2313440121","article-title":"The genetic regulatory architecture and epigenomic basis for age-related changes in rattlesnake venom","volume":"121","author":"Hogan","year":"2024","journal-title":"P Natl Acad Sci USA"},{"key":"2026060909570848800_evag131-B41","doi-asserted-by":"publisher","first-page":"10667","DOI":"10.1073\/pnas.0401720101","article-title":"Pattern of polymorphism after strong artificial selection in a domestication event","volume":"101","author":"Innan","year":"2004","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2026060909570848800_evag131-B42","first-page":"191","article-title":"Notes on the natural history of the Caicos Islands Dwarf Boa, Tropidophis greenwayi","volume":"22","author":"Iverson","year":"1986","journal-title":"Caribb J Sci"},{"key":"2026060909570848800_evag131-B43","doi-asserted-by":"publisher","first-page":"5281","DOI":"10.1038\/ncomms6281","article-title":"On the unfounded enthusiasm for soft selective sweeps","volume":"5","author":"Jensen","year":"2014","journal-title":"Nat Commun"},{"key":"2026060909570848800_evag131-B44","doi-asserted-by":"publisher","first-page":"1236","DOI":"10.1093\/bioinformatics\/btu031","article-title":"Interproscan 5: genome-scale protein function classification","volume":"30","author":"Jones","year":"2014","journal-title":"Bioinformatics"},{"key":"2026060909570848800_evag131-B45","doi-asserted-by":"publisher","first-page":"5258","DOI":"10.1111\/mec.12933","article-title":"Ontogenetic stage-specific quantitative trait loci contribute to divergence in developmental trajectories of sexually dimorphic fins between medaka populations","volume":"23","author":"Kawajiri","year":"2014","journal-title":"Mol Ecol"},{"key":"2026060909570848800_evag131-B46","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1111\/jzo.12066","article-title":"Evolution on a desert island: body size divergence between the reptiles of Nevada\u2019s Anaho Island and the mainland around Pyramid Lake","volume":"291","author":"Keehn","year":"2013","journal-title":"J Zool"},{"key":"2026060909570848800_evag131-B47","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1111\/j.0014-3820.2005.tb00909.x","article-title":"Rapid and repeated origin of insular gigantism and dwarfism in Australian tiger snakes","volume":"59","author":"Keogh","year":"2005","journal-title":"Evolution"},{"key":"2026060909570848800_evag131-B48","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1038\/s41559-019-0812-7","article-title":"Antarctic blackfin icefish genome reveals adaptations to extreme environments","volume":"3","author":"Kim","year":"2019","journal-title":"Nat Ecol Evol"},{"key":"2026060909570848800_evag131-B49","article-title":"TrimGalore!: a wrapper around Cutadapt and FastQC to consistently apply adapter and quality trimming to FastQ files, with extra functionality for RRBS data","author":"Krueger","year":"2015","journal-title":"Babraham Institute"},{"key":"2026060909570848800_evag131-B50","doi-asserted-by":"publisher","first-page":"1571","DOI":"10.1093\/bioinformatics\/btr167","article-title":"Bismark: a flexible aligner and methylation caller for Bisulfite-seq applications","volume":"27","author":"Krueger","year":"2011","journal-title":"Bioinformatics"},{"key":"2026060909570848800_evag131-B51","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1016\/0092-8674(94)90290-9","article-title":"Hox genes in vertebrate development","volume":"78","author":"Krumlauf","year":"1994","journal-title":"Cell"},{"key":"2026060909570848800_evag131-B52","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1038\/nmeth.1923","article-title":"Fast gapped-read alignment with Bowtie 2","volume":"9","author":"Langmead","year":"2012","journal-title":"Nat Methods"},{"key":"2026060909570848800_evag131-B53","doi-asserted-by":"publisher","first-page":"288","DOI":"10.3389\/fgene.2013.00288","article-title":"Sequencing quality assessment tools to enable data-driven informatics for high throughput genomics","volume":"4","author":"Leggett","year":"2013","journal-title":"Front Genet"},{"key":"2026060909570848800_evag131-B54","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1093\/molbev\/msy209","article-title":"Genome-wide regulatory adaptation shapes population-level genomic landscapes in Heliconius","volume":"36","author":"Lewis","year":"2018","journal-title":"Mol Biol Evol"},{"key":"2026060909570848800_evag131-B55","volume-title":"Islands and snakes: isolation and adaptive evolution","author":"Lillywhite","year":"2019"},{"key":"2026060909570848800_evag131-B56","doi-asserted-by":"publisher","first-page":"1683","DOI":"10.1111\/j.1365-2699.2005.01314.x","article-title":"Body size evolution in insular vertebrates: generality of the island rule","volume":"32","author":"Lomolino","year":"2005","journal-title":"J Biogeogr"},{"key":"2026060909570848800_evag131-B57","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1038\/387070a0","article-title":"Adaptive differentiation following experimental island colonization in Anolis lizards","volume":"387","author":"Losos","year":"1997","journal-title":"Nature"},{"key":"2026060909570848800_evag131-B58","doi-asserted-by":"publisher","first-page":"830","DOI":"10.1038\/nature07893","article-title":"Adaptation and diversification on islands","volume":"457","author":"Losos","year":"2009","journal-title":"Nature"},{"key":"2026060909570848800_evag131-B59","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol"},{"key":"2026060909570848800_evag131-B60","doi-asserted-by":"publisher","first-page":"92","DOI":"10.2307\/1445927","article-title":"Venom ontogeny in the Pacific rattlesnakes Crotalus viridis helleri and C. v. oreganus","volume":"1988","author":"Mackessy","year":"1988","journal-title":"Copeia"},{"key":"2026060909570848800_evag131-B61","doi-asserted-by":"publisher","first-page":"1463","DOI":"10.1016\/j.toxicon.2010.02.028","article-title":"Evolutionary trends in venom composition in the Western Rattlesnakes (crotalus viridis sensu lato): toxicity vs. tenderizers","volume":"55","author":"Mackessy","year":"2010","journal-title":"Toxicon"},{"key":"2026060909570848800_evag131-B62","author":"Mackessy"},{"key":"2026060909570848800_evag131-B63","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1643\/HA03-037.1","article-title":"Ontogenetic variation in venom composition and diet of Crotalus oreganus concolor: a case of venom paedomorphosis?","volume":"2003","author":"Mackessy","year":"2003","journal-title":"Copeia"},{"key":"2026060909570848800_evag131-B64","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1016\/j.tig.2017.11.007","article-title":"Reassessing the role of Hox genes during vertebrate development and evolution","volume":"34","author":"Mallo","year":"2018","journal-title":"Trends Genet"},{"key":"2026060909570848800_evag131-B65","doi-asserted-by":"publisher","first-page":"3951","DOI":"10.1242\/dev.068346","article-title":"The regulation of Hox gene expression during animal development","volume":"140","author":"Mallo","year":"2013","journal-title":"Development"},{"key":"2026060909570848800_evag131-B66","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1016\/j.jprot.2013.11.001","article-title":"Linking the transcriptome and proteome to characterize the venom of the eastern diamondback rattlesnake (Crotalus adamanteus)","volume":"96","author":"Margres","year":"2014","journal-title":"J Proteomics"},{"key":"2026060909570848800_evag131-B67","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1534\/genetics.114.172437","article-title":"Contrasting modes and tempos of venom expression evolution in two snake species","volume":"199","author":"Margres","year":"2015","journal-title":"Genetics"},{"key":"2026060909570848800_evag131-B68","doi-asserted-by":"publisher","first-page":"3405","DOI":"10.1111\/mec.13240","article-title":"Phenotypic integration in the feeding system of the eastern diamondback rattlesnake (Crotalus adamanteus)","volume":"24","author":"Margres","year":"2015","journal-title":"Mol Ecol"},{"key":"2026060909570848800_evag131-B69","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1534\/genetics.115.180547","article-title":"Expression differentiation is constrained to low-expression proteins over ecological timescales","volume":"202","author":"Margres","year":"2016","journal-title":"Genetics"},{"key":"2026060909570848800_evag131-B70","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1016\/j.toxicon.2016.05.005","article-title":"Functional characterizations of venom phenotypes in the eastern diamondback rattlesnake (Crotalus adamanteus) and evidence for expression-driven divergence in toxic activities among populations","volume":"119","author":"Margres","year":"2016","journal-title":"Toxicon"},{"key":"2026060909570848800_evag131-B71","doi-asserted-by":"publisher","first-page":"3099","DOI":"10.1093\/molbev\/msx231","article-title":"Quantity, not quality: rapid adaptation in a polygenic trait proceeded exclusively through expression differentiation","volume":"34","author":"Margres","year":"2017","journal-title":"Mol Biol Evol"},{"key":"2026060909570848800_evag131-B72","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1093\/molbev\/msy207","article-title":"Tipping the scales: the migration-selection balance leans toward selection in snake venoms","volume":"36","author":"Margres","year":"2019","journal-title":"Mol Biol Evol"},{"key":"2026060909570848800_evag131-B73","doi-asserted-by":"publisher","first-page":"1569","DOI":"10.1534\/genetics.117.202655","article-title":"Selection to increase expression, not sequence diversity, precedes gene family origin and expansion in rattlesnake venom","volume":"206","author":"Margres","year":"2017","journal-title":"Genetics"},{"key":"2026060909570848800_evag131-B74","doi-asserted-by":"publisher","first-page":"e2014634118","DOI":"10.1073\/pnas.2014634118","article-title":"The Tiger Rattlesnake genome reveals a complex genotype underlying a simple venom phenotype","volume":"118","author":"Margres","year":"2021","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026060909570848800_evag131-B75","doi-asserted-by":"publisher","first-page":"782","DOI":"10.3390\/toxins13110782","article-title":"Varying intensities of introgression obscure incipient venom-associated speciation in the timber rattlesnake (crotalus horridus)","volume":"13","author":"Margres","year":"2021","journal-title":"Toxins (Basel)"},{"key":"2026060909570848800_evag131-B76","doi-asserted-by":"publisher","first-page":"782","DOI":"10.3390\/toxins13110782","article-title":"Varying intensities of introgression obscure incipient venom-associated speciation in the timber rattlesnake (Crotalus horridus)","volume":"13","author":"Margres","year":"2021","journal-title":"Toxins (Basel)"},{"key":"2026060909570848800_evag131-B77","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1159\/000495712","article-title":"Hox genes in reptile development, epigenetic regulation, and teratogenesis","volume":"157","author":"Mart\u00edn-del Campo","year":"2019","journal-title":"Cytogenet Genome Res"},{"key":"2026060909570848800_evag131-B78","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1023\/A:1023866030544","article-title":"Dealing with zeros and missing values in compositional data sets using nonparametric imputation","volume":"35","author":"Martin-Fernandez","year":"2003","journal-title":"Math Geol"},{"key":"2026060909570848800_evag131-B79","volume-title":"Diamonds in the rough, natural history of the eastern diamondback rattlesnake","author":"Means","year":"2017"},{"key":"2026060909570848800_evag131-B80","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1016\/j.tree.2013.08.003","article-title":"Population genomics of rapid adaptation by soft selective sweeps","volume":"28","author":"Messer","year":"2013","journal-title":"Trends Ecol Evol"},{"key":"2026060909570848800_evag131-B81","doi-asserted-by":"publisher","first-page":"e321","DOI":"10.1371\/journal.pbio.0040321","article-title":"Morphological evolution is accelerated among island mammals","volume":"4","author":"Millien","year":"2006","journal-title":"PLoS Biol"},{"key":"2026060909570848800_evag131-B82","doi-asserted-by":"publisher","first-page":"msaf058","DOI":"10.1093\/molbev\/msaf058","article-title":"A segregating structural variant defines novel venom phenotypes in the eastern diamondback rattlesnake","volume":"42","author":"Nachtigall","year":"2025","journal-title":"Mol Biol Evol"},{"key":"2026060909570848800_evag131-B83","author":"Oksanen"},{"key":"2026060909570848800_evag131-B84","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1016\/s0169-5347(01)02167-x","article-title":"The genetics of species differences","volume":"16","author":"Orr","year":"2001","journal-title":"Trends Ecol Evol"},{"key":"2026060909570848800_evag131-B85","doi-asserted-by":"publisher","first-page":"2139","DOI":"10.1534\/genetics.117.201459","article-title":"Cis- and trans-regulatory effects on gene expression in a natural population of Drosophila melanogaster","volume":"206","author":"Osada","year":"2017","journal-title":"Genetics"},{"key":"2026060909570848800_evag131-B86","doi-asserted-by":"publisher","first-page":"1058","DOI":"10.1101\/gr.276251.121","article-title":"Snake venom gene expression is coordinated by novel regulatory architecture and the integration of multiple co-opted vertebrate pathways","volume":"32","author":"Perry","year":"2022","journal-title":"Genome Res"},{"key":"2026060909570848800_evag131-B87","doi-asserted-by":"publisher","first-page":"304","DOI":"10.12688\/f1000research","article-title":"Gff utilities: gffread and gffcompare","volume":"9","author":"Pertea","year":"2020","journal-title":"F1000Res"},{"key":"2026060909570848800_evag131-B88","doi-asserted-by":"publisher","first-page":"290","DOI":"10.1038\/nbt.3122","article-title":"StringTie enables improved reconstruction of a transcriptome from RNA-seq reads","volume":"33","author":"Pertea","year":"2015","journal-title":"Nat Biotechnology"},{"key":"2026060909570848800_evag131-B89","doi-asserted-by":"publisher","author":"Poplin","year":"2018","DOI":"10.1101\/201178"},{"key":"2026060909570848800_evag131-B90","doi-asserted-by":"publisher","first-page":"2943","DOI":"10.1093\/bioinformatics\/btac166","article-title":"Analysing high-throughput sequencing data in Python with HTSeq 2.0","volume":"38","author":"Putri","year":"2022","journal-title":"Bioinformatics"},{"key":"2026060909570848800_evag131-B91","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1086\/705380","article-title":"Eco-evolutionary feedbacks predict the time course of rapid life-history evolution","volume":"194","author":"Reznick","year":"2019","journal-title":"Am Nat"},{"key":"2026060909570848800_evag131-B92","doi-asserted-by":"publisher","first-page":"1211","DOI":"10.1126\/science.1086949","article-title":"Major ecological transitions in wild sunflowers facilitated by hybridization","volume":"301","author":"Rieseberg","year":"2003","journal-title":"Science"},{"key":"2026060909570848800_evag131-B93","doi-asserted-by":"publisher","first-page":"2375","DOI":"10.1534\/g3.115.020578","article-title":"Post-transcriptional mechanisms contribute little to phenotypic variation in snake venoms","volume":"5","author":"Rokyta","year":"2015","journal-title":"G3 (Bethesda)"},{"key":"2026060909570848800_evag131-B94","doi-asserted-by":"publisher","first-page":"e3249","DOI":"10.7717\/peerj.3249","article-title":"The genetics of venom ontogeny in the eastern diamondback rattlesnake (Crotalus adamanteus)","volume":"5","author":"Rokyta","year":"2017","journal-title":"PeerJ"},{"key":"2026060909570848800_evag131-B95","doi-asserted-by":"publisher","first-page":"609","DOI":"10.1042\/bj1210609","article-title":"Studies on ribonucleic acid synthesis in the venom glands of Vipera palaestinae (Ophidia, Reptilia)","volume":"121","author":"Rotenberg","year":"1971","journal-title":"Biochem J"},{"key":"2026060909570848800_evag131-B96","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1002\/dvdy.24482","article-title":"Hox genes in the adult skeleton: Novel functions beyond embryonic development","volume":"246","author":"Rux","year":"2017","journal-title":"Dev Dynam"},{"key":"2026060909570848800_evag131-B97","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1126\/science.aat7244","article-title":"Convergent regulatory evolution and loss of flight in paleognathous birds","volume":"364","author":"Sackton","year":"2019","journal-title":"Science"},{"key":"2026060909570848800_evag131-B98","doi-asserted-by":"publisher","first-page":"e1010121","DOI":"10.1371\/journal.pgen.1010121","article-title":"HiCuT: an efficient and low input method to identify protein-directed chromatin interactions","volume":"18","author":"Sati","year":"2022","journal-title":"PLoS Genet"},{"key":"2026060909570848800_evag131-B99","doi-asserted-by":"publisher","first-page":"666","DOI":"10.3390\/toxins11110666","article-title":"The diversity of venom: the importance of behavior and venom system morphology in understanding its ecology and evolution","volume":"11","author":"Schendel","year":"2019","journal-title":"Toxins (Basel)"},{"key":"2026060909570848800_evag131-B100","doi-asserted-by":"publisher","first-page":"590","DOI":"10.1101\/gr.240952.118","article-title":"The origins and evolution of chromosomes, dosage compensation, and mechanisms underlying venom regulation in snakes","volume":"29","author":"Schield","year":"2019","journal-title":"Genome Res"},{"key":"2026060909570848800_evag131-B101","doi-asserted-by":"publisher","first-page":"1367","DOI":"10.1038\/s41559-022-01829-5","article-title":"The roles of balancing selection and recombination in the evolution of rattlesnake venom","volume":"6","author":"Schield","year":"2022","journal-title":"Nat Ecol Evol"},{"key":"2026060909570848800_evag131-B102","doi-asserted-by":"publisher","first-page":"57320","DOI":"10.3791\/57320","article-title":"Promoter capture hi-c: high-resolution, genome-wide profiling of promoter interactions","volume":"136","author":"Schoenfelder","year":"2018","journal-title":"J Vis Exp"},{"key":"2026060909570848800_evag131-B103","doi-asserted-by":"publisher","first-page":"659","DOI":"10.3390\/toxins12100659","article-title":"Gradual and discrete ontogenetic shifts in rattlesnake venom composition and assessment of hormonal and ecological correlates","volume":"12","author":"Schonour","year":"2020","journal-title":"Toxins (Basel)"},{"key":"2026060909570848800_evag131-B104","doi-asserted-by":"publisher","first-page":"493","DOI":"10.2307\/2389631","article-title":"Why do larger snakes eat larger prey items?","volume":"5","author":"Shine","year":"1991","journal-title":"Funct Ecol"},{"key":"2026060909570848800_evag131-B105","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1186\/s12915-023-01626-x","article-title":"Snakes on a plain: biotic and abiotic factors determine venom compositional variation in a wide-ranging generalist rattlesnake","volume":"21","author":"Smith","year":"2023","journal-title":"BMC Biol"},{"key":"2026060909570848800_evag131-B106","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-017-17765-5","article-title":"Evidence for divergent patterns of local selection driving venom variation in Mojave Rattlesnakes (Crotalus scutulatus)","volume":"8","author":"Strickland","year":"2018","journal-title":"Sci Rep"},{"key":"2026060909570848800_evag131-B107","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13227-020-00171-w","article-title":"Insights into how development and life-history dynamics shape the evolution of venom","volume":"12","author":"Surm","year":"2021","journal-title":"Evodevo"},{"key":"2026060909570848800_evag131-B108","first-page":"341","volume-title":"robCompositions: an R\u2013package for robust statistical analysis of compositional data","author":"Templ","year":"2011"},{"key":"2026060909570848800_evag131-B109","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1139\/cjz-2019-0137","article-title":"Insular dwarfism in female Eastern Hog-nosed Snakes (Heterodon platirhinos; Dipsadidae) on a barrier island","volume":"98","author":"Vanek","year":"2020","journal-title":"Canadian J Zool"},{"key":"2026060909570848800_evag131-B110","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1111\/j.1558-5646.1973.tb05914.x","article-title":"Body size and numbers of plants and animals","volume":"27","author":"Van Valen","year":"1973","journal-title":"Evolution"},{"key":"2026060909570848800_evag131-B111","doi-asserted-by":"publisher","first-page":"14185","DOI":"10.1073\/pnas.94.25.14185","article-title":"Adaptive evolution via a major gene effect: paedomorphosis in the Mexican axolotl","volume":"94","author":"Voss","year":"1997","journal-title":"P Natl Acad Sci USA"},{"key":"2026060909570848800_evag131-B112","doi-asserted-by":"publisher","first-page":"530","DOI":"10.1016\/j.biocon.2012.11.021","article-title":"Life history constraints contribute to the vulnerability of a declining North American rattlesnake","volume":"159","author":"Waldron","year":"2013","journal-title":"Biol Conserv"},{"key":"2026060909570848800_evag131-B113","doi-asserted-by":"publisher","first-page":"1450","DOI":"10.1093\/nar\/gks1339","article-title":"Transcription factor and chromatin features predict genes associated with eQTLs","volume":"41","author":"Wang","year":"2013","journal-title":"Nucleic Acids Res"},{"key":"2026060909570848800_evag131-B114","doi-asserted-by":"publisher","first-page":"1114","DOI":"10.1016\/j.ajhg.2016.03.029","article-title":"Efficient integrative multi-SNP association analysis via deterministic approximation of posteriors","volume":"98","author":"Wen","year":"2016","journal-title":"Am J Hum Genet"},{"key":"2026060909570848800_evag131-B115","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.xinn.2021.100141","article-title":"clusterprofiler 4.0: a universal enrichment tool for interpreting omics data","volume":"2","author":"Wu","year":"2021","journal-title":"Innovation"},{"key":"2026060909570848800_evag131-B116","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1186\/s12915-021-01208-9","article-title":"Dynamic genetic differentiation drives the widespread structural and functional convergent evolution of snake venom proteinaceous toxins","volume":"20","author":"Xie","year":"2022","journal-title":"BMC Biol"},{"key":"2026060909570848800_evag131-B117","doi-asserted-by":"publisher","first-page":"4928","DOI":"10.1038\/s41467-020-18638-8","article-title":"Chromatin accessibility landscape and regulatory network of high-altitude hypoxia adaptation","volume":"11","author":"Xin","year":"2020","journal-title":"Nat Commun"},{"key":"2026060909570848800_evag131-B118","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13059-020-1929-3","article-title":"From reads to insight: a hitchhiker\u2019s guide to ATAC-seq data analysis","volume":"21","author":"Yan","year":"2020","journal-title":"Genome Biol"},{"key":"2026060909570848800_evag131-B119","doi-asserted-by":"publisher","first-page":"100056","DOI":"10.1016\/j.ecochg.2022.100056","article-title":"A Late-Holocene palynological record of coastal ecological change and climate variability from Apalachicola, Florida, USA","volume":"3","author":"Yao","year":"2022","journal-title":"Clim Change Ecol"},{"key":"2026060909570848800_evag131-B120","doi-asserted-by":"publisher","first-page":"2777","DOI":"10.1093\/molbev\/msaa133","article-title":"Venom systems as models for studying the origin and regulation of evolutionary novelties","volume":"37","author":"Zancolli","year":"2020","journal-title":"Mol Biol Evol"},{"key":"2026060909570848800_evag131-B121","doi-asserted-by":"publisher","first-page":"1084","DOI":"10.3390\/genes12071084","article-title":"How important are structural variants for speciation?","volume":"12","author":"Zhang","year":"2021","journal-title":"Genes (Basel)"},{"key":"2026060909570848800_evag131-B122","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/gb-2008-9-9-r137","article-title":"Model-based analysis of ChIP-Seq (MACS)","volume":"9","author":"Zhang","year":"2008","journal-title":"Genome Biol"},{"key":"2026060909570848800_evag131-B123","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1111\/j.1095-8312.2010.01411.x","article-title":"Adaptive significance of food income in European snakes: body size is related to prey energetics","volume":"100","author":"Zuffi","year":"2010","journal-title":"Biol J Linn Soc"}],"container-title":["Genome Biology and Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/gbe\/advance-article-pdf\/doi\/10.1093\/gbe\/evag131\/68434604\/evag131.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/gbe\/article-pdf\/18\/6\/evag131\/68434604\/evag131.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/gbe\/article-pdf\/18\/6\/evag131\/68434604\/evag131.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,9]],"date-time":"2026-06-09T13:57:19Z","timestamp":1781013439000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/gbe\/article\/doi\/10.1093\/gbe\/evag131\/8698343"}},"subtitle":[],"editor":[{"given":"Matthew","family":"Webster","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,5,29]]},"references-count":123,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2026,6,2]]}},"URL":"https:\/\/doi.org\/10.1093\/gbe\/evag131","relation":{},"ISSN":["1759-6653"],"issn-type":[{"value":"1759-6653","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,6]]},"published":{"date-parts":[[2026,5,29]]},"article-number":"evag131"}}