{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T14:04:58Z","timestamp":1753884298792,"version":"3.41.2"},"reference-count":44,"publisher":"The Company of Biologists","issue":"9","license":[{"start":{"date-parts":[[2025,4,30]],"date-time":"2025-04-30T00:00:00Z","timestamp":1745971200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100004375","name":"Tel Aviv University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100004375","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,5,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>Some duck species mostly swim on the water surface while others frequently dive underwater. We compared the paddling kinematics of mandarin ducks (Axis galericulata) that feed on the surface and diving ferruginous pochards (Aythya nyroca) that feed underwater. Both species were trained to perform the same horizontal, submerged swimming at 1\u2005m depth in a controlled set-up. Mandarins used alternate foot paddling exclusively, while pochards varied their gait between alternate foot paddling and simultaneous paddling with both feet. Unlike mandarins, pochards swam with their body tilted at an angle that was negatively correlated with the swimming speed and limited their foot motion to a smaller arc. Hydrodynamic modeling revealed that lift generated by the webbed foot provided thrust to propel both duck species forward. However, mandarins' feet generated lift-based upthrust that interfered with the need to counter their buoyancy, while pochards directed the foot lift to provide vertical downthrust against their buoyancy. The relatively subtle differences in foot motion between the two species result in a substantial hydrodynamic effect that may hint at the kinematic changes required when transitioning from surface to submerged swimming in the evolution of foot-propelled diving waterfowl.<\/jats:p>","DOI":"10.1242\/jeb.249274","type":"journal-article","created":{"date-parts":[[2025,3,13]],"date-time":"2025-03-13T09:39:02Z","timestamp":1741858742000},"source":"Crossref","is-referenced-by-count":0,"title":["Underwater paddling kinematics and hydrodynamics in a surface swimming duck versus a diving duck"],"prefix":"10.1242","volume":"228","author":[{"given":"Hagar","family":"Csillag","sequence":"first","affiliation":[{"name":"School of Zoology 1 , Faculty of Life Sciences , , Tel Aviv 6997801 ,","place":["Israel"]},{"name":"Tel Aviv University 1 , Faculty of Life Sciences , , Tel Aviv 6997801 ,","place":["Israel"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6267-5471","authenticated-orcid":false,"given":"Gal","family":"Ribak","sequence":"additional","affiliation":[{"name":"School of Zoology 1 , Faculty of Life Sciences , , Tel Aviv 6997801 ,","place":["Israel"]},{"name":"Tel Aviv University 1 , Faculty of Life Sciences , , Tel Aviv 6997801 ,","place":["Israel"]},{"name":"The Steinhardt Museum of Natural History, Israel National Center for Biodiversity Studies 2 , Tel Aviv 6997801 ,","place":["Israel"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"237","published-online":{"date-parts":[[2025,4,30]]},"reference":[{"key":"2025043005184754800_JEB249274C1","doi-asserted-by":"publisher","first-page":"1567","DOI":"10.1242\/jeb.198.7.1567","article-title":"Hydroplaning by ducklings: overcoming limitations to swimming at the water surface","volume":"198","author":"Aigeldinger","year":"1995","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C2","doi-asserted-by":"publisher","first-page":"1745","DOI":"10.1242\/jeb.204.10.1745","article-title":"Dynamics of mallard (Anas platyrhynchos) gastrocnemius function during swimming versus terrestrial locomotion","volume":"204","author":"Biewener","year":"2001","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C3","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780198743156.001.0001","volume-title":"Animal Locomotion","author":"Biewener","year":"2018"},{"key":"2025043005184754800_JEB249274C4","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1086\/318111","article-title":"Energetic costs of surface swimming and diving of birds","volume":"73","author":"Butler","year":"2000","journal-title":"Physiol. Biochem. Zool."},{"key":"2025043005184754800_JEB249274C5","doi-asserted-by":"publisher","first-page":"jeb168831","DOI":"10.1242\/jeb.168831","article-title":"Foot-propelled swimming kinematics and turning strategies in common loons","volume":"221","author":"Clifton","year":"2018","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C6","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1111\/joa.12710","article-title":"Comparative hindlimb myology of foot-propelled swimming birds","volume":"232","author":"Clifton","year":"2018","journal-title":"J. Anat."},{"key":"2025043005184754800_JEB249274C7","doi-asserted-by":"publisher","first-page":"9380","DOI":"10.1073\/pnas.1304838110","article-title":"High flight costs, but low dive costs, in auks support the biomechanical hypothesis for flightlessness in penguins","volume":"110","author":"Elliott","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025043005184754800_JEB249274C8","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1002\/ar.24396","article-title":"A mechanism for evolutionary transition in swimming mode by mammals","volume-title":"Secondary Adaptation of Tetrapods to Life in Water","author":"Fish","year":"2001"},{"key":"2025043005184754800_JEB249274C9","doi-asserted-by":"publisher","first-page":"3974","DOI":"10.1242\/jeb.02482","article-title":"Regulation of stroke pattern and swim speed across a range of current velocities: diving by common eiders wintering in polynyas in the Canadian Arctic","volume":"209","author":"Heath","year":"2006","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C10","doi-asserted-by":"publisher","first-page":"034001","DOI":"10.1088\/1748-3182\/3\/3\/034001","article-title":"Software techniques for two- and three-dimensional kinematic measurements of biological and biomimetic systems","volume":"3","author":"Hedrick","year":"2008","journal-title":"Bioinspir. Biomim."},{"volume-title":"Fluid-Dynamic Lift","year":"1985","author":"Hoerner","key":"2025043005184754800_JEB249274C11"},{"key":"2025043005184754800_JEB249274C12","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1242\/jeb.205.3.371","article-title":"Kinematics of diving Atlantic puffins (Fratercula arctica L.): evidence for an active upstroke","volume":"205","author":"Johansson","year":"2002","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C13","doi-asserted-by":"publisher","first-page":"582","DOI":"10.1038\/35036689","article-title":"Asymmetric toes aid underwater swimming","volume":"407","author":"Johansson","year":"2000","journal-title":"Nature"},{"key":"2025043005184754800_JEB249274C14","doi-asserted-by":"publisher","first-page":"1687","DOI":"10.1242\/jeb.204.10.1687","article-title":"Lift-based paddling in diving grebe","volume":"204","author":"Johansson","year":"2001","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C15","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1038\/nature01695","article-title":"Delta-wing function of webbed feet gives hydrodynamic lift for swimming propulsion in birds","volume":"424","author":"Johansson","year":"2003","journal-title":"Nature"},{"key":"2025043005184754800_JEB249274C16","volume-title":"Ducks, Geese, and Swans of the World","author":"Johnsgard","year":"2010","edition":"Revised edn."},{"key":"2025043005184754800_JEB249274C17","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1007\/s00360-005-0047-6","article-title":"Substitution of heat from exercise and digestion by ducks diving for mussels at varying depths and temperatures","volume":"176","author":"Kaseloo","year":"2006","journal-title":"J. Comp. Physiol. B"},{"key":"2025043005184754800_JEB249274C18","doi-asserted-by":"publisher","first-page":"111","DOI":"10.5038\/2074-1235.50.2.1475","article-title":"Common mergansers Mergus merganser use wings to pursue a fish underwater","volume":"50","author":"Lapsansky","year":"2022","journal-title":"Mar. Ornithol."},{"key":"2025043005184754800_JEB249274C19","doi-asserted-by":"publisher","first-page":"24302","DOI":"10.1038\/s41598-021-03868-7","article-title":"Mitochondrial genome of the critically endangered Baer's Pochard, Aythya baeri, and its phylogenetic relationship with other Anatidae species","volume":"11","author":"Liu","year":"2021","journal-title":"Sci. Rep."},{"key":"2025043005184754800_JEB249274C20","first-page":"2","article-title":"Mechanics of underwater swimming in foot-propelled diving birds","author":"Lovvorn","year":"1990","journal-title":"Acta XX Congr. Int. Ornithol"},{"key":"2025043005184754800_JEB249274C21","doi-asserted-by":"publisher","first-page":"e03780","DOI":"10.1002\/ecs2.3780","article-title":"Feeding on epibenthic zooplankton by Long-tailed Ducks: patch structure, profitability, and food web implications","volume":"12","author":"Lovvorn","year":"2021","journal-title":"Ecosphere"},{"key":"2025043005184754800_JEB249274C22","doi-asserted-by":"publisher","first-page":"2888","DOI":"10.1139\/z91-407","article-title":"Body mass, volume, and buoyancy of some aquatic birds, and their relation to locomotor strategies","volume":"69","author":"Lovvorn","year":"1991","journal-title":"Can. J. Zool."},{"key":"2025043005184754800_JEB249274C23","doi-asserted-by":"publisher","first-page":"62","DOI":"10.2307\/1352335","article-title":"Biomechanical conflicts between adaptations for diving and aerial flight in estuarine birds","volume":"17","author":"Lovvorn","year":"1994","journal-title":"Estuaries"},{"key":"2025043005184754800_JEB249274C24","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1242\/jeb.159.1.89","article-title":"Mechanics of underwater locomotion in diving ducks: drag, buoyancy and acceleration in a size gradient of species","volume":"159","author":"Lovvorn","year":"1991","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C25","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1002\/jez.1721","article-title":"Walking, paddling, waddling: 3D kinematics Anatidae locomotion (Callonetta leucophrys)","volume":"317","author":"Provini","year":"2012","journal-title":"J. Exp. Zool. Part A Ecol. Genet. Physiol."},{"key":"2025043005184754800_JEB249274C26","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1242\/jeb.118.1.247","article-title":"Three-dimensional reconstruction of animal flight paths and the turning flight of microchiropteran bats","volume":"118","author":"Rayner","year":"1985","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C27","doi-asserted-by":"publisher","first-page":"16217","DOI":"10.1038\/s41598-023-42784-w","article-title":"The hydrodynamic performance of duck feet for submerged swimming resembles oars rather than delta-wings","volume":"13","author":"Ribak","year":"2023","journal-title":"Sci. Rep."},{"key":"2025043005184754800_JEB249274C28","doi-asserted-by":"publisher","first-page":"2101","DOI":"10.1242\/jeb.00997","article-title":"How do cormorants counter buoyancy during submerged swimming?","volume":"207","author":"Ribak","year":"2004","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C29","doi-asserted-by":"publisher","first-page":"e12565","DOI":"10.1371\/journal.pone.0012565","article-title":"Drag-based \u2018hovering\u2019 in ducks: the hydrodynamics and energetic cost of bottom feeding","volume":"5","author":"Ribak","year":"2010","journal-title":"PLoS ONE"},{"key":"2025043005184754800_JEB249274C30","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1007\/s00360-007-0225-9","article-title":"Costs of diving by wing and foot propulsion in a sea duck, the white-winged scoter","volume":"178","author":"Richman","year":"2008","journal-title":"J. Comp. Physiol. B"},{"key":"2025043005184754800_JEB249274C31","doi-asserted-by":"publisher","first-page":"3801","DOI":"10.1002\/ar.25512","article-title":"How does the shape of the wing and hindlimb bones of aquatic birds relate to their locomotor abilities?","volume":"307","author":"Segesdi","year":"2024","journal-title":"Anat. Rec."},{"key":"2025043005184754800_JEB249274C32","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1242\/jeb.190.1.155","article-title":"Diving energetics in Lesser scaup (Aythyta affinis, Eyton)","volume":"190","author":"Stephenson","year":"1994","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C33","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1016\/0034-5687(94)00130-R","article-title":"Respiratory and plumage gas volumes in unrestrained diving ducks (Aythya affinis)","volume":"100","author":"Stephenson","year":"1995","journal-title":"Respir. Physiol."},{"key":"2025043005184754800_JEB249274C34","unstructured":"Taylor-Burt, K. R.\n           (2020). How to waddle with a paddle: a study of duck hindlimb anatomy, kinematics, and muscle function across behaviors and species. PhD thesis, Harvard University."},{"key":"2025043005184754800_JEB249274C35","doi-asserted-by":"publisher","first-page":"jeb223743","DOI":"10.1242\/jeb.223743","article-title":"Aquatic and terrestrial takeoffs require different hindlimb kinematics and muscle function in mallard ducks","volume":"223","author":"Taylor-Burt","year":"2020","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C36","doi-asserted-by":"publisher","first-page":"1843","DOI":"10.1242\/jeb.100529","article-title":"A protocol and calibration method for accurate multi-camera field videography","volume":"217","author":"Theriault","year":"2014","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C37","doi-asserted-by":"publisher","first-page":"234","DOI":"10.2307\/4070795","article-title":"The use of the wings and feet by diving birds","volume":"26","author":"Townsend","year":"1909","journal-title":"Auk"},{"key":"2025043005184754800_JEB249274C38","doi-asserted-by":"publisher","first-page":"20222056","DOI":"10.1098\/rspb.2022.2056","article-title":"Diving into a dead-end: asymmetric evolution of diving drives diversity and disparity shifts in waterbirds","volume":"289","author":"Tyler","year":"2022","journal-title":"Proc. R. Soc. B."},{"volume-title":"Comparative Biomechanics","year":"2013","author":"Vogel","key":"2025043005184754800_JEB249274C39"},{"key":"2025043005184754800_JEB249274C40","doi-asserted-by":"publisher","first-page":"1875","DOI":"10.1098\/rspb.2000.1224","article-title":"Mechanical performance of aquatic rowing and flying","volume":"267","author":"Walker","year":"2000","journal-title":"Proc. R. Soc. B"},{"key":"2025043005184754800_JEB249274C41","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1098\/rspb.2002.2252","article-title":"Stroke and glide of wing-propelled divers: deep diving seabirds adjust surge frequency to buoyancy change with depth","volume":"270","author":"Watanuki","year":"2003","journal-title":"Proc. R. Soc. B"},{"key":"2025043005184754800_JEB249274C42","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1086\/285409","article-title":"Diving birds in cold water: do Archimedes and Boyle determine energetic costs?","volume":"140","author":"Wilson","year":"1992","journal-title":"Am. Nat."},{"key":"2025043005184754800_JEB249274C43","doi-asserted-by":"publisher","first-page":"1085","DOI":"10.1242\/jeb.00208","article-title":"Leg morphology and locomotion in birds: requirements for force and speed during ankle flexion","volume":"206","author":"Zeffer","year":"2003","journal-title":"J. Exp. Biol."},{"key":"2025043005184754800_JEB249274C44","doi-asserted-by":"publisher","first-page":"461","DOI":"10.1046\/j.1095-8312.2003.00200.x","article-title":"Functional correlation between habitat use and leg morphology in birds (Aves)","volume":"79","author":"Zeffer","year":"2003","journal-title":"Biol. J. Linn. Soc."}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.249274\/3639099\/jeb249274.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.249274\/3646626\/jeb249274.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.249274\/3646626\/jeb249274.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,4,30]],"date-time":"2025-04-30T09:18:56Z","timestamp":1746004736000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/228\/9\/jeb249274\/367858\/Underwater-paddling-kinematics-and-hydrodynamics"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,4,30]]},"references-count":44,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2025,5,1]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.249274","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"type":"print","value":"0022-0949"},{"type":"electronic","value":"1477-9145"}],"subject":[],"published":{"date-parts":[[2025,4,30]]},"article-number":"jeb249274"}}