Stroke frequency, but not swimming speed, is related to body size in free-ranging seabirds, pinnipeds and cetaceans.
Open accessAbstract
It is obvious, at least qualitatively, that small animals move their locomotory apparatus faster than large animals: small insects move their wings invisibly fast, while large birds flap their wings slowly. However, quantitative observations have been difficult to obtain from free-ranging swimming animals. We surveyed the swimming behaviour of animals ranging from 0.5 kg seabirds to 30 000 kg sperm whales using animal-borne accelerometers. Dominant stroke cycle frequencies of swimming specialist seabirds and marine mammals were proportional to mass(-0.29) (R(2)= 0.99, n = 17 groups), while propulsive swimming speeds of 1-2 m s(-1) were independent of body size. This scaling relationship, obtained from breath-hold divers expected to swim optimally to conserve oxygen, does not agree with recent theoretical predictions for optimal swimming. Seabirds that use their wings for both swimming and flying stroked at a lower frequency than other swimming specialists of the same size, suggesting a morphological trade-off with wing size and stroke frequency representing a compromise. In contrast, foot-propelled diving birds such as shags had similar stroke frequencies as other swimming specialists. These results suggest that muscle characteristics may constrain swimming during cruising travel, with convergence among diving specialists in the proportions and contraction rates of propulsive muscles.
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Ocean sunfish oscillate the dorsal and anal fins synchronously.
“This pattern of acceleration was previously reported in penguins [7], which flap both flippers synchronously, indicating that ocean sunfish oscillate the dorsal and anal fins synchronously.”
From Functional Dorsoventral Symmetry in Relation to Lift-Based Swimming in the Ocean Sunfish Mola mola (2008) · cites this paper as [7]
The Results report two positive peaks in penguins’ longitudinal acceleration during a flipper stroke cycle, supporting the cited comparison of acceleration patterns. The passage does not state that penguins flap both flippers synchronously. The inference about ocean sunfish is the citing authors’ own, not a finding attributed to this source. (LLM, conf=0.82)
Experimental — accuracy not yet measured
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Cites
Flying and swimming animals cruise at a Strouhal number tuned for high power efficiency
Taylor, G. K., Nudds, R. L., & Thomas, A. L. R. (2003) · Nature · doi:10.1038/nature02000 · Cited by 994 · see citing works
Flagged first, then by citations
Research works printed but not linked: 22
- Akamatsu T Wang D Wang K Wei Z Zhao Q Naito Y Diving behaviour of freshwater finless porpoises (Neophocaena phocaenoides) in an oxbow of the Yangtze River, China ICES J. Mar. Sci 59 2002 438 443 doi:10.1006/jmsc.2001.1159 · DOI
- Alexander R.M Principles of animal locomotion 2003 Princeton University Press Princeton, NJ
- Anderson J.M Streitlien K Barrett D.S Triantafyllou M.S Oscillating foils of high propulsive efficiency J. Fluid Mech 360 1998 41 72 doi:10.1017/S0022112097008392 · DOI
- Bejan A Marden J.H Unifying constructal theory for scale effects in running, swimming and flying J. Exp. Biol 209 2006 238 248 doi:10.1242/jeb.01974 16391346 · DOI
- Hui C.A Penguin swimming. I. Hydrodynamics Physiol. Zool 61 1988 333 343
- Johansson L.C Aldrin B.S.W Kinematics of diving Atlantic puffins (Fratercula arctica L.): evidence for an active upstroke J. Exp. Biol 205 2002 371 378 11854373
- Johnson M.P Tyack P.L A digital acoustic recording tag for measuring the response of wild marine mammals to sound IEEE J. Ocean. Eng 28 2003 3 12 doi:10.1109/JOE.2002.808212 · DOI
- Kawabe R Naito Y Sato K Miyashita K Yamashita N Direct measurement of the swimming speed, tailbeat, and body angle of Japanese flounder (Paralichthys olivaceus) ICES J. Mar. Sci 61 2004 1080 1087 doi:10.1016/j.icesjms.2004.07.014 · DOI
- Miller P.J.O Johnson M.P Tyack P.L Terray E.A Swimming gaits, passive drag and buoyancy of diving sperm whales Physeter macrocephalus J. Exp. Biol 207 2004 1953 1967 doi:10.1242/jeb.00993 15107448 · DOI
- Pennycuick C.J Wingbeat frequency of birds in steady cruising flight: new data and improved predictions J. Exp. Biol 199 1996 1613 1618 9319516
- Rohr J.J Fish F.E Strouhal numbers and optimization of swimming by odontocete cetaceans J. Exp. Biol 207 2004 1633 1642 doi:10.1242/jeb.00948 15073196 · DOI
- Sato K Naito Y Kato A Niizuma Y Watanuki Y Charrassin J.B Bost C.-A Handrich Y LeMaho Y Buoyancy and maximal diving depth in penguins: do they control inhaling air volume? J. Exp. Biol 205 2002 1189 1197 11948196
- Sato K Mitani Y Cameron M.F Siniff D.B Naito Y Factors affecting stroking patterns and body angle in diving Weddell seals under natural conditions J. Exp. Biol 206 2003 1461 1470 doi:10.1242/jeb.00265 12654885 · DOI
- Sato K Charrassin J.B Bost C.-A Naito Y Why do macaroni penguins choose shallow body angles that result in longer descent and ascent durations? J. Exp. Biol 207 2004 4057 4065 doi:10.1242/jeb.01265 15498951 · DOI
- Sato K Ponganis P.J Habara Y Naito Y Emperor penguins adjust swim speed according to the above-water height of ice holes through which they exit J. Exp. Biol 208 2005 2549 2554 doi:10.1242/jeb.01665 15961741 · DOI
- Tanaka H Takagi Y Naito Y Swimming speeds and buoyancy compensation of migrating adult chum salmon Oncorhynchus keta revealed by speed/depth/acceleration data logger J. Exp. Biol 204 2001 3895 3904 11807107
- Videler J.J Nolet B.A Cost of swimming measured at optimum speed: scale effects, differences between swimming styles, taxonomic groups and submerged and surface swimming Comp. Biochem. Physiol 97A 1990 91 99 doi:10.1016/0300-9629(90)90155-L 1982941 · DOI
- Wallace B.P Williams C.L Paladino F.V Morreale S.J Lindstrom R.T Spotila J.R Bioenergetics and diving activity of internesting leatherback turltes Dermochelys coriacea at Parque Nacional Marino Las Baulas, Costa Rica J. Exp. Biol 208 2005 3873 3884 doi:10.1242/jeb.01860 16215215 · DOI
- Watanuki Y Niizuma Y Gabrielsen G.W Sato K Naito Y Stroke and glide of wing-propelled divers: deep diving seabirds adjust surge frequency to buoyancy change with depth Proc. R. Soc. B 270 2003 483 488 doi:10.1098/rspb.2002.2252 PMC1691262 12641902 · DOI
- Watanuki Y Takahashi A Daunt F Wanless S Harris M Sato K Naito Y Regulation of stroke and glide in a foot-propelled avian diver J. Exp. Biol 208 2005 2207 2216 doi:10.1242/jeb.01639 15939764 · DOI
- Watanuki Y Wanless S Harris M Lovvorn J.R Miyazaki M Tanaka H Sato K Swim speed and stroke patterns in wing-propelled divers: a comparison among alcids and a penguin J. Exp. Biol 209 2006 1217 1230 doi:10.1242/jeb.02128 16547294 · DOI
- Williams T.M The evolution of cost efficient swimming in marine mammals: limits to energetic optimization Phil. Trans. R. Soc. B 354 1999 193 201 doi:10.1098/rstb.1999.0371 · DOI
Cited by
Sorted by citations
Metabolic rate and body size are linked with perception of temporal information
Healy, K., McNally, L., Ruxton, G. D., Cooper, N., & Jackson, A. L. (2013) · Animal Behaviour · doi:10.1016/j.anbehav.2013.06.018 · Cited by 202 · see citing works
Cade, D. E., Gough, W. T., Czapanskiy, M. F., Fahlbusch, J. A., Kahane-Rapport, S. R., Linsky, J. M. J., Nichols, R. C., Oestreich, W. K., Wisniewska, D. M., Friedlaender, A. S., et al. (2021) · Animal Biotelemetry · doi:10.1186/s40317-021-00256-w · Cited by 78 · see citing works
Functional Dorsoventral Symmetry in Relation to Lift-Based Swimming in the Ocean Sunfish Mola mola
Watanabe, Y., & Sato, K. (2008) · PLoS ONE · doi:10.1371/journal.pone.0003446 · Cited by 57 · see citing works
“as stroking activities and body inclination, and have been extensively applied to free-ranging marine mammals and seabirds [7].”
BackgroundMethodSelf-citation
- Authors
- Sato, K., Watanuki, Y., Takahashi, A., Miller, P. J. O., Tanaka, H., Kawabe, R., Ponganis, P. J., Handrich, Y., Akamatsu, T., Watanabe, Y., Mitani, Y., Costa, D. P., Bost, C.-A., Aoki, K., Amano, M., Trathan, P., Shapiro, A., & Naito, Y.
- Affiliations
- The University of Tokyo (JP)
- Hokkaido University (JP)
- National Institute of Polar Research (JP)
- University of St Andrews (GB)
- Nagasaki University (JP)
- Scripps Institution of Oceanography (US)
- Centre National de la Recherche Scientifique (FR)
- Japan Fisheries Research and Education Agency (JP)
- Tokyo Institute of Technology (JP)
- University of California, Santa Cruz (US)
- British Antarctic Survey, Natural Environment Research CouncilHigh Cross, Madingley Road, Cambridge CB3 0ET, UK
- Woods Hole Oceanographic Institution (US)
- Corresponding author
- Sato, Katsufumi
- ORCID
- Venue
- Proceedings of the Royal Society B: Biological Sciences
- Publisher
- The Royal Society
- Published
- 2006-12-05
- Volume
- 274
- Issue
- 1609
- Pages
- 471-477
- ISSN
- 0080-4649 (ISSN-L)
- 0962-8452
- 1471-2954
- PMC
- PMC1766382
- OpenAlex
- W2166061105
- Cited by
- 217
- References
- 1
- Citations by year
- 2026: 11 · 2025: 6 · 2024: 8 · 2023: 13 · 2022: 14 · 2021: 12 · 2020: 23 · 2019: 10 · 2018: 10 · 2017: 11 · 2016: 6 · 2015: 9 · 2014: 7 · 2013: 12 · 2012: 9
- FWCI
- 8.92
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Locations
- Proceedings of the Royal Society B Biological Sciences (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- HAL (submittedVersion)
- Open Access Server of the Woods Hole Scientific Community (Woods Hole Scientific Community) (submittedVersion, cc-by)
- NERC Open Research Archive (Natural Environment Research Council) (submittedVersion)
- Funders
- National Science Foundation
- University of Tokyo
- Natural Environment Research Council (bas010017)
- Japan Society for the Promotion of Science
- Topics
- Physical Sciences › Environmental Science › Ecology › Marine animal studies overview
- Physical Sciences › Engineering › Aerospace Engineering › Biomimetic flight and propulsion mechanisms
- Physical Sciences › Environmental Science › Ecology › Physiological and biochemical adaptations
- Keywords
- stroke frequency, swimming speed, body size scaling, seabirds, pinnipeds, cetaceans, diving birds
- MeSH terms
- Animals, Biomechanical Phenomena, Birds, Cetacea, Caniformia, Swimming, Body Size
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review