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They possess up to three visual pigments: a short\u2010wavelength\u2010sensitive opsin (SWS1), a medium\/long\u2010wavelength\u2010sensitive opsin (LWS), and rhodopsin (RH1). Nocturnal snakes have duplex retinas containing both rod and cone photoreceptors, whereas diurnal caenophidian (\u201cadvanced\u201d) snakes exhibit simplex \u201call\u2010cone\u201d retinas, lacking morphologically typical rods. In this study, we analyzed photoreceptor morphology in the retinas of caenophidian snakes using high\u2010resolution scanning electron microscopy (SEM) and examined visual\u2010opsin expression patterns with immunohistochemistry (IHC). Our analyses revealed remarkable interspecific variability in visual\u2010cell morphology. Light microscopy showed that in all sampled diurnal caenophidians, photoreceptors expressing RH1 exhibit a gross cone\u2010like morphology. However, SEM analysis revealed a subset of photoreceptors with distinct features\u2014thinner inner segments and rod\u2010like synaptic terminals\u2014suggesting they are transmuted, cone\u2010like rods. In retinal sections from nocturnal caenophidian snakes, coexpression of the cone opsins SWS1 and LWS in individual cones was observed, whereas rhodopsin expression remained restricted to morphologically typical rods and showed no coexpression. In contrast, diurnal caenophidians commonly coexpress rhodopsin and SWS1 in single cones, with some instances of triple coexpression (SWS1, RH1, and LWS) in single cones. We evaluated the patterns of spatial distribution of RH1\u2010 and SWS1\u2010expressing photoreceptors, as well as SWS1\u00a0+\u00a0RH1 multiopsin cones, in wholemounted retinas of ten species. Our findings revealed considerable species\u2010specific variation in photoreceptor density, topography, and opsin coexpression patterns. IHC results suggest that in some species, rhodopsin is not only expressed in transmuted, cone\u2010like rods but may also be co\u2010opted by UV\/violet\u2010sensitive (SWS1\u2010expressing) cones. These findings underscore the exceptional diversity and adaptive innovation in snake visual systems. The unique features and striking interspecific differences in their photoreceptors highlight snakes as an outstanding taxon for studying vertebrate visual\u2010system function and evolution.<\/jats:p>","DOI":"10.1002\/cne.70092","type":"journal-article","created":{"date-parts":[[2025,10,7]],"date-time":"2025-10-07T10:02:30Z","timestamp":1759831350000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Exceptional Visual\u2010Opsin Coexpression and Phenotypic Diversity in Outer\u2010Retinal Photoreceptors of Caenophidian Snakes"],"prefix":"10.1002","volume":"533","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1561-2769","authenticated-orcid":false,"given":"Einat","family":"Hauzman","sequence":"first","affiliation":[{"name":"Department of Experimental Psychology, Psychology Institute University of S\u00e3o Paulo  S\u00e3o Paulo Brazil"},{"name":"Natural History Museum  London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-7820-1119","authenticated-orcid":false,"given":"Silke","family":"Haverkamp","sequence":"additional","affiliation":[{"name":"Department of Computational Neuroethology Max Planck Institute for Neurobiology of Behavior  Bonn Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5272-1713","authenticated-orcid":false,"given":"Juliana H.","family":"Tashiro","sequence":"additional","affiliation":[{"name":"Department of Experimental Psychology, Psychology Institute University of S\u00e3o Paulo  S\u00e3o Paulo Brazil"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5475-9210","authenticated-orcid":false,"given":"Irene L.","family":"G\u00fcgel","sequence":"additional","affiliation":[{"name":"Department of Computational Neuroethology Max Planck Institute for Neurobiology of Behavior  Bonn Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6691-3951","authenticated-orcid":false,"given":"Natalia F.","family":"Torello\u2010Viera","sequence":"additional","affiliation":[{"name":"Department of Experimental Psychology, Psychology Institute University of S\u00e3o Paulo  S\u00e3o Paulo Brazil"},{"name":"Institute of Biosciences Federal University of Mato Grosso  Cuiab\u00e1 Brazil"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3318-7193","authenticated-orcid":false,"given":"Tha\u00eds B.","family":"Guedes","sequence":"additional","affiliation":[{"name":"Department of Biodiversity, Bioscience Institute S\u00e3o Paulo State University Julio de Mesquita Filho UNESP  Rio Claro Brazil"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0277-0239","authenticated-orcid":false,"given":"Pavel","family":"N\u011bmec","sequence":"additional","affiliation":[{"name":"Department of Zoology, Faculty of Science Charles University  Prague Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8035-4719","authenticated-orcid":false,"given":"Nicholas R.","family":"Casewell","sequence":"additional","affiliation":[{"name":"Centre for Snakebite Research &amp; 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