{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:54:23Z","timestamp":1782867263179,"version":"3.54.5"},"reference-count":56,"publisher":"The Royal Society","issue":"2074","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"The University of Western Australia Collaborative Grant Program"},{"name":"Smithsonian Institution Fellowship Program"},{"DOI":"10.13039\/100000008","name":"David and Lucile Packard Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000008","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Rathbun Endowment for Crustacean Research"},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Kenneth Jay Boss Fellowship Program"},{"name":"Emmy Noether Junior Research Group Grant"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,7,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>In the largest habitat on earth\u2014the deep open ocean\u2014water depth is typically linked to adaptations for camouflage and vision. Recently, it was hypothesized that association behaviour is a primary evolutionary driver of the great eye diversity in a group of crustaceans, the hyperiid amphipods (suborder Hyperiidea). Here, we test that hypothesis by investigating the link between association behaviour (symbiotic or free swimming), depth, eye complexity and camouflage across a broad range of hyperiid diversity. We collated a unique dataset containing observations of 48 of the 80 hyperiid genera from a comprehensive literature review, blackwater scuba photographs and nearly 30 years of submersible observations. By scoring association behaviour in 5759 in situ observations, we show that approximately one third of hyperiid genera have adopted a free-swimming lifestyle. This supports a shift from the current assumption that all hyperiids live in close association with gelatinous animals, to one allowing a free-swimming, predatory lifestyle in many genera. Using phylogenetic generalized linear mixed model (PGLMM) analyses, we show that a free-swimming lifestyle correlates with body transparency and more complex eyes at upper mesopelagic depths. Our results support the hypothesis that a free-swimming lifestyle drives eye complexity while increasing the need for transparency.<\/jats:p>","DOI":"10.1098\/rspb.2025.2837","type":"journal-article","created":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:11:37Z","timestamp":1782864697000},"source":"Crossref","is-referenced-by-count":0,"title":["<i>In situ<\/i>\n                    observations reveal a link between association behaviour, camouflage and eye complexity in midwater amphipods"],"prefix":"10.1098","volume":"293","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9403-257X","authenticated-orcid":false,"given":"Vanessa I.","family":"Stenvers","sequence":"first","affiliation":[{"name":"Smithsonian National Museum of Natural History, 1","place":["Washington,, DC, USA, 20013,"],"department":["Department of Invertebrate Zoology,"]},{"name":"GEOMAR Helmholtz Centre for Ocean Research Kiel, 2","place":["Kiel,, Schleswig-Holstein, Germany, 24148,"],"department":["Marine Ecology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4629-9362","authenticated-orcid":false,"given":"Jan M.","family":"Hemmi","sequence":"additional","affiliation":[{"name":"Smithsonian National Museum of Natural History, 1","place":["Washington,, DC, USA, 20013,"],"department":["Department of Invertebrate Zoology,"]},{"name":"University of Western Australia, 3","place":["Perth,, Western Australia, Australia, 6009,"],"department":["School of Biological Schiences,"]},{"name":"University of Western Australia, 4","place":["Perth,, Western Australia, Australia, 6009,"],"department":["UWA Oceans Institute,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kyra L.","family":"Schlining","sequence":"additional","affiliation":[{"name":"Monterey Bay Aquarium Research Institute, 5","place":["Moss Landing,, CA, USA, 95039,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6225-444X","authenticated-orcid":false,"given":"Richard","family":"Collins","sequence":"additional","affiliation":[{"name":"Florida Museum of Natural History, 6","place":["Gainesville,, FL, USA, 32611-2710,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Linda","family":"Ianniello","sequence":"additional","affiliation":[{"name":"Unaffiliated, 7"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Holly F.","family":"Waters","sequence":"additional","affiliation":[{"name":"Smithsonian National Museum of Natural History, 1","place":["Washington,, DC, USA, 20013,"],"department":["Department of Invertebrate Zoology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0305-1159","authenticated-orcid":false,"given":"C. Anela","family":"Choy","sequence":"additional","affiliation":[{"name":"University of California San Diego, 8","place":["La Jolla,, CA, USA, 92037,"],"department":["Scripps Institution of Oceanography,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven H. D.","family":"Haddock","sequence":"additional","affiliation":[{"name":"Monterey Bay Aquarium Research Institute, 5","place":["Moss Landing,, CA, USA, 95039,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1784-0466","authenticated-orcid":false,"given":"Bruce H.","family":"Robison","sequence":"additional","affiliation":[{"name":"Monterey Bay Aquarium Research Institute, 5","place":["Moss Landing,, CA, USA, 95039,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4330-6507","authenticated-orcid":false,"given":"Henk-Jan","family":"Hoving","sequence":"additional","affiliation":[{"name":"GEOMAR Helmholtz Centre for Ocean Research Kiel, 2","place":["Kiel,, Schleswig-Holstein, Germany, 24148,"],"department":["Marine Ecology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4226-9257","authenticated-orcid":false,"given":"Karen J.","family":"Osborn","sequence":"additional","affiliation":[{"name":"Smithsonian National Museum of Natural History, 1","place":["Washington,, DC, USA, 20013,"],"department":["Department of Invertebrate Zoology,"]},{"name":"University of Western Australia, 3","place":["Perth,, Western Australia, Australia, 6009,"],"department":["School of Biological Schiences,"]},{"name":"University of Western Australia, 4","place":["Perth,, Western Australia, Australia, 6009,"],"department":["UWA Oceans Institute,"]},{"name":"Monterey Bay Aquarium Research Institute, 5","place":["Moss Landing,, CA, USA, 95039,"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,7,1]]},"reference":[{"key":"2026063020112564800_R1","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/j.jembe.2004.01.012","article-title":"Deep pelagic biology","volume":"300","author":"Robison","year":"2004","journal-title":"J. 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Pap."},{"key":"2026063020112564800_R11","first-page":"1","volume-title":"Annales de l\u2019Institut oc\u00e9anographique monaco, nouvelle s\u00e9rie","author":"Pirlot","year":"1932"},{"key":"2026063020112564800_R12","volume-title":"Adjuncta est materia medica kahirina atque tabula maris rubri geographica","author":"Risso","year":"1816"},{"key":"2026063020112564800_R13","first-page":"1","article-title":"Recherches sur les animaux inf\u00e9rieurs du golfe de Marseille. Deuxi\u00e9me m\u00e9moire: Description des crustac\u00e9s amphipodes parasites des salpes","volume":"17","author":"Marion","year":"1874","journal-title":"Ann. Sci. Nat."},{"key":"2026063020112564800_R14","first-page":"1","article-title":"The hyperiid amphipod, Hyperia galba, a true ecto-parasite on jelly-fish","volume":"1959","author":"Dahl","year":"1959","journal-title":"Universitet i Bergen, Arbok 1959. Naturvidenskapeling Rekke"},{"key":"2026063020112564800_R15","doi-asserted-by":"crossref","first-page":"141","DOI":"10.2307\/1350747","article-title":"Notes on associations between hyperiid amphipods and medusae in Chesapeake and Narragansett Bays and the Niantic River","volume":"4","author":"Bowman","year":"1963","journal-title":"Chesapeake Sci."},{"key":"2026063020112564800_R16","first-page":"11","article-title":"Hyperiid amphipods as crustacean parasitoids associated with gelatinous zooplankton","volume":"18","author":"Laval","year":"1980","journal-title":"Oceanogr. Mar. Biol. Annu. 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Associations with Salpidae","volume":"24","author":"Madin","year":"1977","journal-title":"Deep Sea Res."},{"key":"2026063020112564800_R20","first-page":"632","volume-title":"Hyperiid amphipods (Amphipoda, Hyperiidea) of the world oceans","author":"Vinogradov","year":"1996"},{"key":"2026063020112564800_R21","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1007\/BF00616747","article-title":"The eyes of hyperiid amphipods: relations of optical structure to depth","volume":"164","author":"Land","year":"1989","journal-title":"J. Comp. Physiol."},{"key":"2026063020112564800_R22","doi-asserted-by":"crossref","first-page":"103610","DOI":"10.1016\/j.dsr.2021.103610","article-title":"Extraordinary eyes reveal hidden diversity within the holopelagic genus Paraphronima (Amphipoda: Hyperiidea)","volume":"177","author":"Stenvers","year":"2021","journal-title":"Deep Sea Res. Part I: Oceanographic Res. Pap"},{"key":"2026063020112564800_R23","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1016\/j.cub.2014.12.010","article-title":"A unique apposition compound eye in the mesopelagic hyperiid amphipod Paraphronima gracilis","volume":"25","author":"Baldwin Fergus","year":"2015","journal-title":"Curr. Biol."},{"key":"2026063020112564800_R24","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1010545","article-title":"A new computational model illuminates the extraordinary eyes of Phronima","volume":"18","author":"Bagheri","year":"2022","journal-title":"PLoS Comput. Biol."},{"key":"2026063020112564800_R25","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1017\/S0025315400018981","article-title":"The vertical distribution and diurnal migration of the Crustacea Amphipoda collected during the SOND Cruise, 1965 II. The Hyperiidea and general discussion","volume":"56","author":"Thurston","year":"1976","journal-title":"J. Mar. Biol. Ass."},{"key":"2026063020112564800_R26","first-page":"1","article-title":"MBARI\u2019s video annotation and reference system","author":"Schlining","year":"2006","journal-title":"OCEANS"},{"key":"2026063020112564800_R27","doi-asserted-by":"crossref","DOI":"10.17815\/jlsrf-3-157","article-title":"Manned submersible JAGO","volume":"3","author":"Hissmann","year":"2017","journal-title":"JLSRF"},{"key":"2026063020112564800_R28","article-title":"Data from: In situ observations reveal a link between association behaviour, camouflage and eye complexity in midwater amphipods. Figshare.","author":"Stenvers","year":"2026"},{"key":"2026063020112564800_R29","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1016\/0022-0981(78)90008-4","article-title":"The barrel of the pelagic amphipod Phronima sedentaria (Forsk.) (Crustacea: hyperiidea)","volume":"33","author":"Laval","year":"1978","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"2026063020112564800_R30","doi-asserted-by":"crossref","first-page":"1435","DOI":"10.1017\/S0025315405012610","article-title":"Still alive? Fine structure of the barrels made by Phronima (Crustacea: Amphipoda)","volume":"85","author":"Hirose","year":"2005","journal-title":"J. Mar. Biol. Ass."},{"key":"2026063020112564800_R31","unstructured":"R Core Team\n          . 2022R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing."},{"key":"2026063020112564800_R32","unstructured":"Kolde\n              R\n            \n          . 2019pheatmap: Pretty Heatmaps [software]. Version 1.0.12. Comprehensive R Archive Network (CRAN). 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Biol."},{"key":"2026063020112564800_R36","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.ympev.2012.12.021","article-title":"Molecular phylogenetic evidence for the reorganization of the Hyperiid amphipods, a diverse group of pelagic crustaceans","volume":"67","author":"Hurt","year":"2013","journal-title":"Mol. Phylogenet. Evol."},{"key":"2026063020112564800_R37","doi-asserted-by":"crossref","first-page":"106664","DOI":"10.1016\/j.ympev.2019.106664","article-title":"The late blooming amphipods: global change promoted post-Jurassic ecological radiation despite Palaeozoic origin","volume":"143","author":"Copila\u015f-Ciocianu","year":"2020","journal-title":"Mol. Phylogenet. Evol."},{"key":"2026063020112564800_R38","unstructured":"Biancani\n              LM\n            \n          . 2019Multi-locus phylogenetic analysis of Amphipoda indicates a single origin of the pelagic suborder Hyperiidea. Digital Repository at the University of Maryland. 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Ass."},{"key":"2026063020112564800_R42","first-page":"229","article-title":"Beobachtungen zu Entwicklung und Verhalten von Phronima sedentaria (Forskal), (Amphipoda)","volume":"10","author":"Richter","year":"1978","journal-title":"Senckenbergiana Maritima"},{"key":"2026063020112564800_R43","doi-asserted-by":"crossref","first-page":"79","DOI":"10.2307\/1548433","article-title":"Observations on the anatomy and behavior of Phronima sedentaria (Forskal) (amphipoda: Hyperiidea)","volume":"8","author":"Diebel","year":"1988","journal-title":"J. Crust. Biol."},{"key":"2026063020112564800_R44","doi-asserted-by":"crossref","DOI":"10.1002\/ecy.4410","article-title":"The association of argonauts with gelatinous plankton and other substrates","volume":"105","author":"Villanueva","year":"2024","journal-title":"Ecology"},{"key":"2026063020112564800_R45","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2017.2116","article-title":"Deep pelagic food web structure as revealed by in situ feeding observations","volume":"284","author":"Choy","year":"2017","journal-title":"Proc. R. Soc. B"},{"key":"2026063020112564800_R46","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1093\/icesjms\/fsz010","article-title":"Enlightening the ocean\u2019s twilight zone","volume":"76","author":"Kaartvedt","year":"2019","journal-title":"ICES J. Mar. 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Biol."},{"key":"2026063020112564800_R50","doi-asserted-by":"crossref","first-page":"301","DOI":"10.2307\/1543609","article-title":"Hidden in plain sight: the ecology and physiology of organismal transparency","volume":"201","author":"Johnsen","year":"2001","journal-title":"Biol. Bull."},{"key":"2026063020112564800_R51","doi-asserted-by":"crossref","DOI":"10.1038\/srep45750","article-title":"Quantification of bioluminescence from the surface to the deep sea demonstrates its predominance as an ecological trait","volume":"7","author":"Martini","year":"2017","journal-title":"Sci. Rep."},{"key":"2026063020112564800_R52","doi-asserted-by":"crossref","first-page":"3470","DOI":"10.1016\/j.cub.2020.06.044","article-title":"Ultra-black camouflage in deep-sea fishes","volume":"30","author":"Davis","year":"2020","journal-title":"Curr. Biol."},{"key":"2026063020112564800_R53","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1017\/S1464793103006420","article-title":"Vision in the deep sea","volume":"79","author":"Warrant","year":"2004","journal-title":"Biol. Rev. Camb. Philos. Soc."},{"key":"2026063020112564800_R54","doi-asserted-by":"crossref","first-page":"1147","DOI":"10.1098\/rstb.2000.0656","article-title":"On the functions of double eyes in midwater animals","volume":"355","author":"Land","year":"2000","journal-title":"Phil. Trans. R. Soc. Lond. B"},{"key":"2026063020112564800_R55","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1016\/S0022-0981(02)00533-6","article-title":"Evidence for long-range chemoreceptive tracking of food odour in deep-sea scavengers by scanning sonar data","volume":"285\u2013286","author":"Premke","year":"2003","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"2026063020112564800_R56","article-title":"Supplementary material from: In situ observations reveal a link between association behaviour, camouflage and eye complexity in midwater amphipods. Figshare.","author":"Stenvers","year":"2026"}],"container-title":["Proceedings of the Royal Society B: Biological Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article-pdf\/doi\/10.1098\/rspb.2025.2837\/6149604\/rspb.2025.2837.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article-pdf\/doi\/10.1098\/rspb.2025.2837\/6149604\/rspb.2025.2837.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:11:53Z","timestamp":1782864713000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article\/293\/2074\/20252837\/482372\/In-situ-observations-reveal-a-link-between"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,1]]},"references-count":56,"journal-issue":{"issue":"2074","published-print":{"date-parts":[[2026,7,1]]}},"URL":"https:\/\/doi.org\/10.1098\/rspb.2025.2837","relation":{},"ISSN":["1471-2954"],"issn-type":[{"value":"1471-2954","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,1]]},"article-number":"20252837"}}