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To understand the underlying mechanisms and their evolutionary relationships, we must define each mode and identify its phenotypic and molecular markers. In this study, we focus on a widely dispersed migration mode characterized by dynamic actin-filled pseudopods that we call \u201c\u03b1-motility.\u201d Mining genomic data reveals a clear trend: only organisms with both WASP and SCAR\/WAVE\u2014activators of branched actin assembly\u2014make actin-filled pseudopods. Although SCAR has been shown to drive pseudopod formation, WASP\u2019s role in this process is controversial. We hypothesize that these genes collectively represent a genetic signature of \u03b1-motility because both are used for pseudopod formation. WASP depletion from human neutrophils confirms that both proteins are involved in explosive actin polymerization, pseudopod formation, and cell migration. WASP and WAVE also colocalize to dynamic signaling structures. Moreover, retention of WASP together with SCAR correctly predicts \u03b1-motility in disease-causing chytrid fungi, which we show crawl at &amp;gt;30 \u00b5m\/min with actin-filled pseudopods. By focusing on one migration mode in many eukaryotes, we identify a genetic marker of pseudopod formation, the morphological feature of \u03b1-motility, providing evidence for a widely distributed mode of cell crawling with a single evolutionary origin.<\/jats:p>","DOI":"10.1083\/jcb.201701074","type":"journal-article","created":{"date-parts":[[2017,5,4]],"date-time":"2017-05-04T10:40:20Z","timestamp":1493894420000},"page":"1673-1688","update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":134,"title":["WASP and SCAR are evolutionarily conserved in actin-filled pseudopod-based motility"],"prefix":"10.1083","volume":"216","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9237-9403","authenticated-orcid":true,"given":"Lillian K.","family":"Fritz-Laylin","sequence":"first","affiliation":[{"name":"Department of Cellular and Molecular Pharmacology, Howard Hughes Medical Institute, University of California, San Francisco, San Francisco, CA 94143"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2785-989X","authenticated-orcid":true,"given":"Samuel J.","family":"Lord","sequence":"additional","affiliation":[{"name":"Department of Cellular and Molecular Pharmacology, Howard Hughes Medical Institute, University of California, San Francisco, San Francisco, CA 94143"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0871-5479","authenticated-orcid":true,"given":"R. 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