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Antagonistic coevolutionary relationships are often not simple interactions between a single predator and prey but rather a complex web of interactions between multiple coexisting species. Coevolution between venomous rattlesnakes and small mammals has led to physiological venom resistance in several mammalian taxa. In general, viperid venoms contain large quantities of snake venom metalloproteinase toxins (SVMPs), which are inactivated by SVMP inhibitors expressed in resistant mammals. We explored variation in venom chemistry, SVMP expression, and SVMP resistance across four co\u2010distributed species (California Ground Squirrels, Bryant's Woodrats, Southern Pacific Rattlesnakes, and Red Diamond Rattlesnakes) collected from four different populations in Southern California. Our aim was to understand phenotypic and functional variation in venom and venom resistance in order to compare coevolutionary dynamics of a system involving two sympatric predator\u2013prey pairs to past studies that have focused on single pairs. Proteomic analysis of venoms indicated that these rattlesnakes express different phenotypes when in sympatry, with Red Diamonds expressing more typical viperid venom (with a diversity of SVMPs) and Southern Pacifics expressing a more atypical venom with a broader range of non\u2010enzymatic toxins. We also found that although blood sera from both mammals were generally able to inhibit SVMPs from both rattlesnake species, inhibition depended strongly on the snake population, with snakes from one geographic site expressing SVMPs to which few mammals were resistant. Additionally, we found that Red Diamond venom, rather than woodrat resistance, was locally adapted. Our findings highlight the complexity of coevolutionary relationships between multiple predators and prey that exhibit similar offensive and defensive strategies in sympatry.<\/jats:p>","DOI":"10.1111\/jeb.13907","type":"journal-article","created":{"date-parts":[[2021,7,29]],"date-time":"2021-07-29T04:00:20Z","timestamp":1627531220000},"page":"1447-1465","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":32,"title":["Phenotypic and functional variation in venom and venom resistance of two sympatric rattlesnakes and their prey"],"prefix":"10.1093","volume":"34","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6501-066X","authenticated-orcid":false,"given":"Kelly E.","family":"Robinson","sequence":"first","affiliation":[{"name":"Department of Biology San Diego State University San Diego CA USA"},{"name":"Department of Biology University of Nevada Reno NV USA"},{"name":"Program in Ecology, Evolution and Conservation Biology University of Nevada Reno NV USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3477-3012","authenticated-orcid":false,"given":"Matthew L.","family":"Holding","sequence":"additional","affiliation":[{"name":"Department of Natural Resources and Environmental Science University of Nevada Reno NV USA"},{"name":"Department of Biological Science Florida State University Tallahassee FL USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2516-2607","authenticated-orcid":false,"given":"Malachi D.","family":"Whitford","sequence":"additional","affiliation":[{"name":"Department of Biology San Diego State University San Diego CA USA"},{"name":"Ecology Graduate Group University of California Davis CA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6890-512X","authenticated-orcid":false,"given":"Anthony J.","family":"Saviola","sequence":"additional","affiliation":[{"name":"Department of Molecular Medicine and Neurobiology The Scripps Research Institute La Jolla CA USA"},{"name":"Department of Biochemistry and Molecular Genetics University of Colorado Anschutz Medical Campus Aurora CO USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5267-1672","authenticated-orcid":false,"suffix":"III","given":"John R.","family":"Yates","sequence":"additional","affiliation":[{"name":"Department of Molecular Medicine and Neurobiology The Scripps Research Institute La Jolla CA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1100-1856","authenticated-orcid":false,"given":"Rulon W.","family":"Clark","sequence":"additional","affiliation":[{"name":"Department of Biology San Diego State University San Diego CA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2021,8,9]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/0040-5809(86)90007-9"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ecolsys.31.1.79"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.97.8.4106"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1086\/692110"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2012.1112"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2009.0048"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v067.i01"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1086\/303213"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.0014-3820.2001.tb01293.x"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1086\/376580"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0109651"},{"key":"e_1_2_9_13_1","first-page":"557","volume-title":"The biology of rattlesnakes","author":"Biardi J. 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