{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T23:26:54Z","timestamp":1780615614153,"version":"3.54.1"},"reference-count":112,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,1,1]],"date-time":"2026-01-01T00:00:00Z","timestamp":1767225600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,1,1]],"date-time":"2026-01-01T00:00:00Z","timestamp":1767225600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,10,24]],"date-time":"2025-10-24T00:00:00Z","timestamp":1761264000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000923","name":"Australian Research Council","doi-asserted-by":"publisher","award":["DP190100304"],"award-info":[{"award-number":["DP190100304"]}],"id":[{"id":"10.13039\/501100000923","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Biochimie"],"published-print":{"date-parts":[[2026,1]]},"DOI":"10.1016\/j.biochi.2025.10.010","type":"journal-article","created":{"date-parts":[[2025,10,18]],"date-time":"2025-10-18T22:34:40Z","timestamp":1760826880000},"page":"137-152","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":3,"special_numbering":"C","title":["Deadly innovations: Molecular phylogenetics and evolution of phospholipase A2 toxins in viperid snake venoms"],"prefix":"10.1016","volume":"240","author":[{"given":"Lorenzo","family":"Seneci","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6521-8850","authenticated-orcid":false,"given":"Vivek","family":"Suranse","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2230-8183","authenticated-orcid":false,"given":"Marco","family":"Mancuso","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tobias","family":"Senoner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2965-9461","authenticated-orcid":false,"given":"Bing","family":"Xie","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9073-4758","authenticated-orcid":false,"given":"Ivan","family":"Koludarov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kartik","family":"Sunagar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6661-1283","authenticated-orcid":false,"given":"Bryan G.","family":"Fry","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.biochi.2025.10.010_bib1","first-page":"431","article-title":"Dietary correlates of the origin and radiation of snakes","volume":"23","author":"Greene","year":"1983","journal-title":"Integr. Comp. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib2","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1046\/j.1096-3642.2003.00052.x","article-title":"The evolution of venom-delivery systems in snakes","volume":"137","author":"Jackson","year":"2003","journal-title":"Zool. J. Linn. Soc."},{"key":"10.1016\/j.biochi.2025.10.010_bib3","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1074\/mcp.M700094-MCP200","article-title":"Evolution of an arsenal: structural and functional diversification of the venom system in the advanced snakes (Caenophidia)","volume":"7","author":"Fry","year":"2008","journal-title":"Mol. Cell. Proteomics"},{"key":"10.1016\/j.biochi.2025.10.010_bib4","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1146\/annurev.genom.9.081307.164356","article-title":"The toxicogenomic multiverse: convergent recruitment of proteins into animal venoms","volume":"10","author":"Fry","year":"2009","journal-title":"Annu. Rev. Genomics Hum. Genet."},{"key":"10.1016\/j.biochi.2025.10.010_bib5","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1016\/j.tree.2012.10.020","article-title":"Complex cocktails: the evolutionary novelty of venoms","volume":"28","author":"Casewell","year":"2013","journal-title":"Trends Ecol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib6","series-title":"Handbook of Venoms and Toxins of Reptiles","first-page":"389","article-title":"Snake venom phospholipase A2 toxins","author":"Lomonte","year":"2021"},{"key":"10.1016\/j.biochi.2025.10.010_bib7","article-title":"Life finds a way: reconstructing the evolutionary history of a diverse gene family reveals complexity at the genetic origin of novelty","author":"Koludarov","year":"2019","journal-title":"bioRxiv"},{"key":"10.1016\/j.biochi.2025.10.010_bib8","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1101\/gr.3228405","article-title":"From genome to \"venome\": molecular origin and evolution of the snake venom proteome inferred from phylogenetic analysis of toxin sequences and related body proteins","volume":"15","author":"Fry","year":"2005","journal-title":"Genome Res."},{"key":"10.1016\/j.biochi.2025.10.010_bib9","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S1388-1981(00)00105-0","article-title":"The expanding superfamily of phospholipase A2 enzymes: classification and characterization","volume":"1488","author":"Six","year":"2000","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"10.1016\/j.biochi.2025.10.010_bib10","series-title":"Venomous Reptiles and their Toxins: Evolution, Pathophysiology, and Biodiscovery","first-page":"335","article-title":"Group II phospholipase A2 enzymes","author":"Sunagar","year":"2015"},{"key":"10.1016\/j.biochi.2025.10.010_bib11","doi-asserted-by":"crossref","first-page":"2897","DOI":"10.1007\/s00018-008-8113-3","article-title":"Cellular pathology induced by snake venom phospholipase A2 myotoxins and neurotoxins: common aspects of their mechanisms of action","volume":"65","author":"Montecucco","year":"2008","journal-title":"Cell. Mol. Life Sci."},{"key":"10.1016\/j.biochi.2025.10.010_bib12","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.toxicon.2013.06.019","article-title":"Predicting function from sequence in a large multifunctional toxin family","volume":"72","author":"Malhotra","year":"2013","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib13","doi-asserted-by":"crossref","DOI":"10.3390\/toxins8090248","article-title":"Varespladib (LY315920) appears to be a potent, broad-spectrum, inhibitor of snake venom phospholipase A2 and a possible pre-referral treatment for envenomation","volume":"8","author":"Lewin","year":"2016","journal-title":"Toxins"},{"key":"10.1016\/j.biochi.2025.10.010_bib14","doi-asserted-by":"crossref","DOI":"10.1016\/j.toxicon.2023.107024","article-title":"Lys49 myotoxins, secreted phospholipase A2-like proteins of viperid venoms: a comprehensive review","volume":"224","author":"Lomonte","year":"2023","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib15","doi-asserted-by":"crossref","first-page":"11627","DOI":"10.1016\/S0021-9258(17)39077-4","article-title":"A comparison of the crystal structure of phospholipase A2 from bovine pancreas and Crotalus atrox venom","volume":"260","author":"Renetseder","year":"1985","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.biochi.2025.10.010_bib16","doi-asserted-by":"crossref","first-page":"5364","DOI":"10.1021\/acs.jmedchem.3c00097","article-title":"Catalytically active snake venom PLA2 enzymes: an overview of its elusive mechanisms of reaction","volume":"66","author":"Castro-Amorim","year":"2023","journal-title":"J. Med. Chem."},{"key":"10.1016\/j.biochi.2025.10.010_bib17","doi-asserted-by":"crossref","DOI":"10.3390\/toxins16020071","article-title":"Viper venom phospholipase A2 database: the structural and functional anatomy of a primary toxin in envenomation","volume":"16","author":"de Oliveira","year":"2024","journal-title":"Toxins"},{"key":"10.1016\/j.biochi.2025.10.010_bib18","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1111\/j.1432-1033.1980.tb04982.x","article-title":"Correlation of enzymatic activity and anticoagulant properties of phospholipase A2","volume":"112","author":"Verheij","year":"1980","journal-title":"Eur. J. Biochem."},{"key":"10.1016\/j.biochi.2025.10.010_bib19","doi-asserted-by":"crossref","first-page":"1587","DOI":"10.1016\/S0041-0101(01)00141-6","article-title":"Identification of the myotoxic site of the Lys49 phospholipase A2 from Agkistrodon piscivorus piscivorus snake venom: synthetic C-terminal peptides from Lys49, but not from Asp49 myotoxins, exert membrane-damaging activities","volume":"39","author":"N\u00faez","year":"2001","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib20","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1016\/j.toxicon.2009.10.013","article-title":"The C-terminal region of a Lys49 myotoxin mediates Ca2+ influx in C2C12 myotubes","volume":"55","author":"Cintra-Francischinelli","year":"2010","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib21","doi-asserted-by":"crossref","first-page":"1229","DOI":"10.1016\/S0041-0101(96)00112-2","article-title":"Accelerated evolution of snake venom phospholipase A2 isozymes for acquisition of diverse physiological functions","volume":"34","author":"Ogawa","year":"1996","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib22","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1007\/PL00006450","article-title":"Accelerated evolution and molecular surface of venom phospholipase A2 enzymes","volume":"48","author":"Kini","year":"1999","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib23","doi-asserted-by":"crossref","DOI":"10.1186\/1471-2148-7-2","article-title":"Inventing an arsenal: adaptive evolution and neofunctionalization of snake venom phospholipase A2 genes","volume":"7","author":"Lynch","year":"2007","journal-title":"BMC Evol. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib24","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1007\/s00239-008-9067-7","article-title":"Rapid evolution by positive selection and gene gain and loss: PLA2 venom genes in closely related Sistrurus rattlesnakes with divergent diets","volume":"66","author":"Gibbs","year":"2008","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib25","doi-asserted-by":"crossref","DOI":"10.3390\/toxins14060420","article-title":"Contextual constraints: dynamic evolution of snake venom phospholipase A2","volume":"14","author":"Suranse","year":"2022","journal-title":"Toxins"},{"key":"10.1016\/j.biochi.2025.10.010_bib26","doi-asserted-by":"crossref","DOI":"10.1093\/gigascience\/giac024","article-title":"The rise of genomics in snake venom research: recent advances and future perspectives","volume":"11","author":"Rao","year":"2022","journal-title":"GigaScience"},{"key":"10.1016\/j.biochi.2025.10.010_bib27","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1093\/jhered\/esae026","article-title":"Whole snake genomes from eighteen families of snakes (Serpentes: caenophidia) and their applications to systematics","volume":"115","author":"Roberts","year":"2024","journal-title":"J. Hered."},{"key":"10.1016\/j.biochi.2025.10.010_bib28","doi-asserted-by":"crossref","first-page":"2959","DOI":"10.1016\/j.cell.2023.05.030","article-title":"Large-scale snake genome analyses provide insights into vertebrate development","volume":"186","author":"Peng","year":"2023","journal-title":"Cell"},{"key":"10.1016\/j.biochi.2025.10.010_bib29","doi-asserted-by":"crossref","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. USA"},{"key":"10.1016\/j.biochi.2025.10.010_bib30","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1016\/j.toxicon.2015.08.009","article-title":"The importance of being genomic: non-coding and coding sequences suggest different models of toxin multi-gene family evolution","volume":"107","author":"Malhotra","year":"2015","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib31","doi-asserted-by":"crossref","DOI":"10.3389\/fphar.2020.574925","article-title":"How the toxin got its toxicity","volume":"11","author":"Jackson","year":"2020","journal-title":"Front. Pharmacol."},{"key":"10.1016\/j.biochi.2025.10.010_bib32","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1038\/nature06151","article-title":"Gene duplication and the adaptive evolution of a classic genetic switch","volume":"449","author":"Hittinger","year":"2007","journal-title":"Nature"},{"key":"10.1016\/j.biochi.2025.10.010_bib33","doi-asserted-by":"crossref","first-page":"762","DOI":"10.1038\/nature07092","article-title":"Escape from adaptive conflict after duplication in an anthocyanin pathway gene","volume":"454","author":"Des Marais","year":"2008","journal-title":"Nature"},{"key":"10.1016\/j.biochi.2025.10.010_bib34","doi-asserted-by":"crossref","first-page":"21593","DOI":"10.1073\/pnas.1007883107","article-title":"Evolution of an antifreeze protein by neofunctionalization under escape from adaptive conflict","volume":"107","author":"Deng","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biochi.2025.10.010_bib35","doi-asserted-by":"crossref","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":"10.1016\/j.biochi.2025.10.010_bib36","doi-asserted-by":"crossref","first-page":"2553","DOI":"10.1093\/molbev\/msu236","article-title":"ExaBayes: massively parallel bayesian tree inference for the whole-genome era","volume":"31","author":"Aberer","year":"2014","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib37","doi-asserted-by":"crossref","first-page":"1530","DOI":"10.1093\/molbev\/msaa015","article-title":"IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era","volume":"37","author":"Minh","year":"2020","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib38","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1093\/molbev\/msx281","article-title":"UFBoot2: improving the ultrafast bootstrap approximation","volume":"35","author":"Hoang","year":"2017","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib39","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1093\/nar\/gkae268","article-title":"Interactive Tree of Life (iTOL) v6: recent updates to the phylogenetic tree display and annotation tool","volume":"52","author":"Letunic","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.biochi.2025.10.010_bib40","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1093\/sysbio\/sys029","article-title":"MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space","volume":"61","author":"Ronquist","year":"2012","journal-title":"Syst. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib41","doi-asserted-by":"crossref","first-page":"1586","DOI":"10.1093\/molbev\/msm088","article-title":"Paml 4: phylogenetic analysis by maximum likelihood","volume":"24","author":"Yang","year":"2007","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib42","doi-asserted-by":"crossref","first-page":"1107","DOI":"10.1093\/molbev\/msi097","article-title":"Bayes empirical Bayes inference of amino acid sites under positive selection","volume":"22","author":"Yang","year":"2005","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib43","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1002764","article-title":"Detecting individual sites subject to episodic diversifying selection","volume":"8","author":"Murrell","year":"2012","journal-title":"PLoS Genet."},{"key":"10.1016\/j.biochi.2025.10.010_bib44","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1093\/bioinformatics\/bti079","article-title":"HyPhy: hypothesis testing using phylogenies","volume":"21","author":"Pond","year":"2005","journal-title":"Bioinformatics"},{"key":"10.1016\/j.biochi.2025.10.010_bib45","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1093\/molbev\/mst030","article-title":"FUBAR: a fast, unconstrained bayesian approximation for inferring selection","volume":"30","author":"Murrell","year":"2013","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib46","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1093\/nar\/gkw408","article-title":"ConSurf 2016: an improved methodology to estimate and visualize evolutionary conservation in macromolecules","volume":"44","author":"Ashkenazy","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.biochi.2025.10.010_bib47","doi-asserted-by":"crossref","DOI":"10.1002\/pro.4582","article-title":"Using evolutionary data to make sense of macromolecules with a \"face-lifted\" ConSurf","volume":"32","author":"Yariv","year":"2023","journal-title":"Protein Sci."},{"key":"10.1016\/j.biochi.2025.10.010_bib48","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with AlphaFold","volume":"596","author":"Jumper","year":"2021","journal-title":"Nature"},{"key":"10.1016\/j.biochi.2025.10.010_bib49","series-title":"Pymol: an Open-Source Molecular Graphics Tool, CCP4 Newsletter on Protein Crystallography","author":"DeLano","year":"2002"},{"key":"10.1016\/j.biochi.2025.10.010_bib50","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1002\/(SICI)1096-987X(199802)19:3<319::AID-JCC6>3.0.CO;2-W","article-title":"Exact and efficient analytical calculation of the accessible surface areas and their gradients for macromolecules","volume":"19","author":"Fraczkiewicz","year":"1998","journal-title":"J. Comput. Chem."},{"key":"10.1016\/j.biochi.2025.10.010_bib51","doi-asserted-by":"crossref","first-page":"7112","DOI":"10.1109\/TPAMI.2021.3095381","article-title":"ProtTrans: toward understanding the language of life through self-supervised learning","volume":"44","author":"Elnaggar","year":"2022","journal-title":"IEEE Trans. Pattern Anal. Mach. Intell."},{"key":"10.1016\/j.biochi.2025.10.010_bib52","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1002\/(SICI)1097-0266(199606)17:6<441::AID-SMJ819>3.0.CO;2-G","article-title":"The application of cluster analysis in strategic management research: an analysis and critique","volume":"17","author":"Ketchen","year":"1996","journal-title":"Strateg. Manag. J."},{"key":"10.1016\/j.biochi.2025.10.010_bib53","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/0377-0427(87)90125-7","article-title":"Silhouettes: a graphical aid to the interpretation and validation of cluster analysis","volume":"20","author":"Rousseeuw","year":"1987","journal-title":"J. Comput. Appl. Math."},{"key":"10.1016\/j.biochi.2025.10.010_bib54","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0210236","article-title":"Clustering algorithms: a comparative approach","volume":"14","author":"Rodriguez","year":"2019","journal-title":"PLoS One"},{"key":"10.1016\/j.biochi.2025.10.010_bib55","doi-asserted-by":"crossref","DOI":"10.1016\/j.jmb.2025.168940","article-title":"ProtSpace: a tool for visualizing protein space","volume":"437","author":"Senoner","year":"2025","journal-title":"J. Mol. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib56","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1038\/nbt.4314","article-title":"Dimensionality reduction for visualizing single-cell data using UMAP","volume":"37","author":"Becht","year":"2018","journal-title":"Nat. Biotechnol."},{"key":"10.1016\/j.biochi.2025.10.010_bib57","first-page":"437","article-title":"Purification and properties of two phospholipase A2 enzymes from berg adder (Bitis atropos) venom","volume":"97","author":"van Zyl","year":"2001","journal-title":"South Afr. J. Sci."},{"key":"10.1016\/j.biochi.2025.10.010_bib58","doi-asserted-by":"crossref","first-page":"4861","DOI":"10.1038\/s41467-023-40550-0","article-title":"Domain loss enabled evolution of novel functions in the snake three-finger toxin gene superfamily","volume":"14","author":"Koludarov","year":"2023","journal-title":"Nat. Commun."},{"key":"10.1016\/j.biochi.2025.10.010_bib59","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1042\/BJ20040125","article-title":"Molecular evolution and structure-function relationships of crotoxin-like and asparagine-6-containing phospholipases A2 in pit viper venoms","volume":"381","author":"Chen","year":"2004","journal-title":"Biochem. J."},{"key":"10.1016\/j.biochi.2025.10.010_bib60","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1016\/j.jmb.2011.07.027","article-title":"Crystal structure of crotoxin reveals key residues involved in the stability and toxicity of this potent heterodimeric \u03b2-neurotoxin","volume":"412","author":"Faure","year":"2011","journal-title":"J. Mol. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib61","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1005596","article-title":"The rise and fall of an evolutionary innovation: contrasting strategies of venom evolution in ancient and young animals","volume":"11","author":"Sunagar","year":"2015","journal-title":"PLoS Genet."},{"key":"10.1016\/j.biochi.2025.10.010_bib62","doi-asserted-by":"crossref","first-page":"2434","DOI":"10.1016\/j.cub.2016.07.038","article-title":"The deep origin and recent loss of venom toxin genes in rattlesnakes","volume":"26","author":"Dowell","year":"2016","journal-title":"Curr. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib63","doi-asserted-by":"crossref","first-page":"462","DOI":"10.1016\/j.bbagen.2005.11.022","article-title":"N49 phospholipase A2, a unique subgroup of snake venom group II phospholipase A2","volume":"1760","author":"Wei","year":"2006","journal-title":"Biochim. Biophys. Acta Gen. Subj."},{"key":"10.1016\/j.biochi.2025.10.010_bib64","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.toxicon.2007.08.007","article-title":"Systemic and local myotoxicity induced by snake venom group II phospholipases A2: comparison between crotoxin, crotoxin B and a Lys49 PLA2 homologue","volume":"51","author":"Guti\u00e9rrez","year":"2008","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib65","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0109846","article-title":"Synergism between basic Asp49 and Lys49 phospholipase A2 myotoxins of viperid snake venom in vitro and in vivo","volume":"9","author":"Mora-Obando","year":"2014","journal-title":"PLoS One"},{"key":"10.1016\/j.biochi.2025.10.010_bib66","doi-asserted-by":"crossref","first-page":"1758","DOI":"10.1016\/j.jprot.2010.06.001","article-title":"Snake venomics and antivenomics of Crotalus durissus subspecies from Brazil: assessment of geographic variation and its implication on snakebite management","volume":"73","author":"Boldrini-Fran\u00e7a","year":"2010","journal-title":"J. Proteonomics"},{"key":"10.1016\/j.biochi.2025.10.010_bib67","doi-asserted-by":"crossref","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. Proteome Res."},{"key":"10.1016\/j.biochi.2025.10.010_bib68","doi-asserted-by":"crossref","first-page":"6684","DOI":"10.1046\/j.1432-1327.2000.01766.x","article-title":"Phospholipases A2 from Callosellasma rhodostoma venom gland. Cloning and sequencing of 10 of the cDNAs, three-dimensional modelling and chemical modification of the major isozyme","volume":"267","author":"Tsai","year":"2000","journal-title":"Eur. J. Biochem."},{"key":"10.1016\/j.biochi.2025.10.010_bib69","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.ympev.2016.07.029","article-title":"Diversification in vipers: phylogenetic relationships, time of divergence and shifts in speciation rates","volume":"105","author":"Alencar","year":"2016","journal-title":"Mol. Phylogenet. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib70","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1016\/j.toxicon.2006.06.021","article-title":"Discovery of novel [arg49]phospholipase A2 isozymes from Protobothrops elegans venom and regional evolution of Crotalinae snake venom phospholipase A2 isozymes in the southwestern islands of Japan and Taiwan","volume":"48","author":"Chijiwa","year":"2006","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib71","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1016\/j.toxicon.2007.11.018","article-title":"Crystal structure of a novel myotoxic Arg49 phospholipase A2 homolog (zhaoermiatoxin) from Zhaoermia mangshanensis snake venom: insights into Arg49 coordination and the role of Lys122 in the polarization of the C-terminus","volume":"51","author":"Murakami","year":"2008","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib72","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1016\/j.biochi.2006.05.003","article-title":"Purification, characterization and cytokine release function of a novel Arg-49 phospholipase A2 from the venom of Protobothrops mucrosquamatus","volume":"88","author":"Wei","year":"2006","journal-title":"Biochimie"},{"key":"10.1016\/j.biochi.2025.10.010_bib73","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1007\/s00239-002-2400-7","article-title":"Interisland evolution of Trimeresurus flavoviridis venom phospholipase A2 isozymes","volume":"56","author":"Chijiwa","year":"2003","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib74","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1017\/S0030605311001037","article-title":"Population status, distribution and conservation needs of the endangered Mangshan pit viper Protobothrops mangshanensis of China","volume":"47","author":"Gong","year":"2013","journal-title":"Oryx"},{"key":"10.1016\/j.biochi.2025.10.010_bib75","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.toxicon.2016.02.014","article-title":"Structures of Azemiops feae venom phospholipases and cys-rich-secretory protein and implications for taxonomy and toxinology","volume":"114","author":"Tsai","year":"2016","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib76","doi-asserted-by":"crossref","DOI":"10.1093\/gbe\/evac082","article-title":"De novo genome assembly highlights the role of lineage-specific gene duplications in the evolution of venom in Fea's viper (Azemiops feae)","volume":"14","author":"Myers","year":"2022","journal-title":"Genome. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib77","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1134\/S1607672912010103","article-title":"The new peptide from the Fea's viper Azemiops feae venom interacts with nicotinic acetylcholine receptors","volume":"442","author":"Utkin","year":"2012","journal-title":"Dokl. Biochem. Biophys."},{"key":"10.1016\/j.biochi.2025.10.010_bib78","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1074\/mcp.M112.023135","article-title":"Differential evolution and neofunctionalization of snake venom metalloprotease domains","volume":"12","author":"Brust","year":"2013","journal-title":"Mol. Cell. Proteomics"},{"key":"10.1016\/j.biochi.2025.10.010_bib79","doi-asserted-by":"crossref","first-page":"2697","DOI":"10.1046\/j.1432-1033.2003.03629.x","article-title":"Sequences and structural organization of phospholipase A2 genes from Vipera aspis aspis, V. aspis zinnikeri and Vipera berus berus venom: identification of the origin of a new viper population based on ammodytin I1 heterogeneity","volume":"270","author":"Guillemin","year":"2003","journal-title":"Eur. J. Biochem."},{"key":"10.1016\/j.biochi.2025.10.010_bib80","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1016\/j.toxicon.2013.05.017","article-title":"Cytotoxic activities of [ser49]phospholipase A2 from the venom of the saw-scaled vipers Echis ocellatus, Echis pyramidum leakeyi, Echis carinatus sochureki, and Echis coloratus","volume":"71","author":"Conlon","year":"2013","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib81","doi-asserted-by":"crossref","first-page":"961","DOI":"10.1042\/bj3190961","volume":"319","author":"Polgar","year":"1996","journal-title":"Biochem. J."},{"key":"10.1016\/j.biochi.2025.10.010_bib82","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1016\/j.ympev.2008.08.019","article-title":"A nesting of vipers: phylogeny and historical biogeography of the Viperidae (Squamata: serpentes)","volume":"49","author":"Wuster","year":"2008","journal-title":"Mol. Phylogenet. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib83","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1016\/j.ympev.2009.08.002","article-title":"When continents collide: phylogeny, historical biogeography and systematics of the medically important viper genus Echis (Squamata: serpentes: viperidae)","volume":"53","author":"Pook","year":"2009","journal-title":"Mol. Phylogenet. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib84","doi-asserted-by":"crossref","DOI":"10.1038\/s41598-020-72871-1","article-title":"Thermal melanism explains macroevolutionary variation of dorsal pigmentation in Eurasian vipers","volume":"10","author":"Mart\u00ednez-Freir\u00eda","year":"2020","journal-title":"Sci. Rep."},{"key":"10.1016\/j.biochi.2025.10.010_bib85","doi-asserted-by":"crossref","first-page":"3425","DOI":"10.1111\/j.1365-294X.2006.03031.x","article-title":"Phylogeography and Pleistocene refugia of the adder (Vipera berus) as inferred from mitochondrial DNA sequence data","volume":"15","author":"Ursenbacher","year":"2006","journal-title":"Mol. Ecol."},{"key":"10.1016\/j.biochi.2025.10.010_bib86","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.toxicon.2009.10.025","article-title":"Understanding the in vitro neuromuscular activity of snake venom Lys49 phospholipase A2 homologues","volume":"55","author":"Gallacci","year":"2010","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib87","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1016\/0041-0101(82)90057-5","article-title":"Site of action of caudoxin, a neurotoxic phospholipase A2 from the horned puff adder (Bitis caudalis) venom","volume":"20","author":"Lee","year":"1982","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib88","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1007\/s12640-021-00330-4","article-title":"Widespread and differential neurotoxicity in venoms from the Bitis genus of viperid snakes","volume":"39","author":"Youngman","year":"2021","journal-title":"Neurotox. Res."},{"key":"10.1016\/j.biochi.2025.10.010_bib89","doi-asserted-by":"crossref","first-page":"1586","DOI":"10.3390\/toxins6051586","article-title":"In vitro toxic effects of puff adder (Bitis arietans) venom, and their neutralization by antivenom","volume":"6","author":"Fernandez","year":"2014","journal-title":"Toxins"},{"key":"10.1016\/j.biochi.2025.10.010_bib90","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1042\/bj3110895","article-title":"Molecular cloning and characterization of a neurotoxic phospholipase A2 from the venom of Taiwan habu (Trimeresurus mucrosquamatus)","volume":"311","author":"Tsai","year":"1995","journal-title":"Biochem. J."},{"key":"10.1016\/j.biochi.2025.10.010_bib91","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1016\/0041-0101(93)90255-H","article-title":"Purification, sequencing and characterization of single amino acid-substituted phospholipase A2 isozymes from Trimeresurus gramineus (green habu snake) venom","volume":"31","author":"Fukagawa","year":"1993","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib92","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1016\/S0014-5793(02)03205-2","article-title":"Toxicity evolution of Vipera aspis aspis venom: identification and molecular modeling of a novel phospholipase A2 heterodimer neurotoxin","volume":"527","author":"Jan","year":"2002","journal-title":"FEBS Lett."},{"key":"10.1016\/j.biochi.2025.10.010_bib93","doi-asserted-by":"crossref","first-page":"722","DOI":"10.1107\/S0907444907016204","article-title":"Structural and pharmacological comparison of daboiatoxin from Daboia russelli siamensis with viperotoxin F and vipoxin from other vipers","volume":"63","author":"Gopalan","year":"2007","journal-title":"Acta. Crystallogr. D Biol. Crystallogr."},{"key":"10.1016\/j.biochi.2025.10.010_bib94","doi-asserted-by":"crossref","first-page":"10911","DOI":"10.1073\/pnas.1920011117","article-title":"The origin and diversification of a novel protein family in venomous snakes","volume":"117","author":"Giorgianni","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biochi.2025.10.010_bib95","doi-asserted-by":"crossref","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":"10.1016\/j.biochi.2025.10.010_bib96","series-title":"Snakes: Morphology, Function, and Ecology","first-page":"343","article-title":"Evolutionary and clinical implications of the snake venom system: dynamic diversification","author":"Fry","year":"2023"},{"key":"10.1016\/j.biochi.2025.10.010_bib97","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1146\/annurev.genet.39.073003.112240","article-title":"Concerted and birth-and-death evolution of multigene families","volume":"39","author":"Nei","year":"2005","journal-title":"Annu. Rev. Genet."},{"key":"10.1016\/j.biochi.2025.10.010_bib98","doi-asserted-by":"crossref","first-page":"2172","DOI":"10.3390\/toxins5112172","article-title":"Three-fingered RAVERs: rapid accumulation of variations in exposed residues of snake venom toxins","volume":"5","author":"Sunagar","year":"2013","journal-title":"Toxins"},{"key":"10.1016\/j.biochi.2025.10.010_bib99","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1098\/rspb.1979.0086","article-title":"The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme","volume":"21","author":"Gould","year":"1979","journal-title":"Proc. Roy. Soc. Lond. B"},{"key":"10.1016\/j.biochi.2025.10.010_bib100","doi-asserted-by":"crossref","first-page":"2994","DOI":"10.1093\/bioinformatics\/bts548","article-title":"DLRS: gene tree evolution in light of a species tree","volume":"28","author":"Sjostrand","year":"2012","journal-title":"Bioinformatics"},{"key":"10.1016\/j.biochi.2025.10.010_bib101","doi-asserted-by":"crossref","first-page":"1147","DOI":"10.1016\/j.toxicon.2005.02.018","article-title":"Structure-function relationships and mechanism of anticoagulant phospholipase A2 enzymes from snake venoms","volume":"45","author":"Kini","year":"2005","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib102","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/j.toxlet.2020.11.010","article-title":"Anticoagulant Micrurus venoms: targets and neutralization","volume":"337","author":"Dashevsky","year":"2021","journal-title":"Toxicol. Lett."},{"key":"10.1016\/j.biochi.2025.10.010_bib103","doi-asserted-by":"crossref","DOI":"10.3390\/toxins8110309","article-title":"Rapid radiations and the race to redundancy: an investigation of the evolution of Australian elapid snake venoms","volume":"8","author":"Jackson","year":"2016","journal-title":"Toxins"},{"key":"10.1016\/j.biochi.2025.10.010_bib104","doi-asserted-by":"crossref","first-page":"885","DOI":"10.1016\/j.toxicon.2003.11.008","article-title":"An overview of lysine-49 phospholipase A2 myotoxins from crotalid snake venoms and their structural determinants of myotoxic action","volume":"42","author":"Lomonte","year":"2003","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib105","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1016\/j.toxicon.2015.01.006","article-title":"Neuromuscular effects of venoms and crotoxin-like proteins from Crotalus durissus ruruima and Crotalus durissus cumanensis","volume":"96","author":"Cavalcante","year":"2015","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib106","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/j.jprot.2014.09.009","article-title":"Structures and functions of crotoxin-like heterodimers and acidic phospholipases A2 from Gloydius intermedius venom: insights into the origin of neurotoxic-type rattlesnakes","volume":"112","author":"Yang","year":"2015","journal-title":"J. Proteonomics"},{"key":"10.1016\/j.biochi.2025.10.010_bib107","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1111\/j.1432-1033.1983.tb07860.x","article-title":"Classification of phospholipases A2 according to sequence. Evolutionary and pharmacological implications","volume":"137","author":"Dufton","year":"1983","journal-title":"Eur. J. Biochem."},{"key":"10.1016\/j.biochi.2025.10.010_bib108","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1016\/j.toxicon.2017.10.026","article-title":"Coevolution takes the sting out of it: evolutionary biology and mechanisms of toxin resistance in animals","volume":"140","author":"Arbuckle","year":"2017","journal-title":"Toxicon"},{"key":"10.1016\/j.biochi.2025.10.010_bib109","doi-asserted-by":"crossref","first-page":"1513","DOI":"10.1111\/jeb.13347","article-title":"Local prey community composition and genetic distance predict venom divergence among populations of the northern Pacific rattlesnake (Crotalus oreganus)","volume":"31","author":"Holding","year":"2018","journal-title":"J. Evol. Biol."},{"key":"10.1016\/j.biochi.2025.10.010_bib110","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1007\/s00239-003-2461-2","article-title":"Molecular evolution and phylogeny of elapid snake venom three-finger toxins","volume":"57","author":"Fry","year":"2003","journal-title":"J. Mol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib111","doi-asserted-by":"crossref","first-page":"2391","DOI":"10.1093\/molbev\/msn179","article-title":"Evolution of snake venom disintegrins by positive Darwinian selection","volume":"25","author":"Ju\u00e1rez","year":"2008","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.biochi.2025.10.010_bib112","doi-asserted-by":"crossref","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."}],"container-title":["Biochimie"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0300908425002342?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0300908425002342?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,3,19]],"date-time":"2026-03-19T09:03:28Z","timestamp":1773911008000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0300908425002342"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1]]},"references-count":112,"alternative-id":["S0300908425002342"],"URL":"https:\/\/doi.org\/10.1016\/j.biochi.2025.10.010","relation":{},"ISSN":["0300-9084"],"issn-type":[{"value":"0300-9084","type":"print"}],"subject":[],"published":{"date-parts":[[2026,1]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Deadly innovations: Molecular phylogenetics and evolution of phospholipase A2 toxins in viperid snake venoms","name":"articletitle","label":"Article Title"},{"value":"Biochimie","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.biochi.2025.10.010","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2025 The Authors. Published by Elsevier B.V.","name":"copyright","label":"Copyright"}]}}