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Katsufumi Sato

Publications and source records attributed to Katsufumi Sato.

9 recordsLinked to original sources

Individual variation in feeding habitat use by adult female green sea turtles (Chelonia mydas): are they obligately neritic herbivores?

Satellite telemetry and stable isotope analysis were used to confirm that oceanic areas (where water depths are >200 m) are alternative feeding habitats for adult female green sea turtles (Chelonia mydas), which have been thought to be obligate herbivores in neritic areas (where depths are <200 m). Four females were tagged with satellite transmitters and tracked during post-nesting periods from Ogasawara Islands, Japan. Three females migrated to neritic habitats, while transmissions from another female ceased in an oceanic habitat. The overall mean nighttime dive depths during oceanic swimming periods in two females were <20 m, implying that the main function of their nighttime dives were resting with neutral buoyancy, whereas the means in two other females were >20 m, implying that they not only rested, but also foraged on macroplankton that exhibit diel vertical migration. Comparisons of stable carbon and nitrogen isotope ratios between 89 females and the prey items in a three-source mixing model estimated that 69% of the females nesting on Ogasawara Islands mainly used neritic habitats and 31% mainly used oceanic habitats. Out of four females tracked by satellite, two females were inferred from isotope ratios to be neritic herbivores and the two others oceanic planktivores. Although post-nesting movements for four females were not completely consistent with the inferences from isotope ratios, possibly due to short tracking periods (28-42 days), their diving behaviors were consistent with the inferences. There were no relationships between body size and the two isotope ratios, indicating a lack of size-related differences in feeding habitat use by adult female green turtles, which was in contrast with loggerhead sea turtles (Caretta caretta). These results and previous findings suggest that ontogenetic habitat shifts by sea turtles are facultative, and consequently, their life histories are polymorphic.

Animal Migration↗

Swim speeds and stroke patterns in wing-propelled divers: a comparison among alcids and a penguin.

In diving birds, the volume and resulting buoyancy of air spaces changes with dive depth, and hydrodynamic drag varies with swim speed. These factors are important in the dive patterns and locomotion of alcids that use their wings both for aerial flight and underwater swimming and of penguins that use their wings only for swimming. Using small data-loggers on free-ranging birds diving to 20-30 m depth, we measured depth at 1 Hz and surge and heave accelerations at 32-64 Hz of four species of alcids (0.6-1.0 kg mass) and the smallest penguin species (1.2 kg). Low- and high-frequency components of the fluctuation of acceleration yielded estimates of body angles and stroke frequencies, respectively. Swim speed was estimated from body angle and rate of depth change. Brünnich's (Uria lomvia) and common (Uria aalge) guillemots descended almost vertically, whereas descent of razorbills (Alca torda), rhinoceros auklets (Cerorhinca monocerata) and little penguins (Eudyptula minor) was more oblique. For all species, swim speed during descent was within a relatively narrow range. Above depths of 20-30 m, where they were all positively buoyant, all species ascended without wing stroking. During descent, little penguins made forward accelerations on both the upstroke and downstroke regardless of dive depth. By contrast, descending alcids produced forward accelerations on both upstroke and downstroke at depths of <10 m but mainly on the downstroke at greater depths; this change seemed to correspond to the decrease of buoyancy with increasing depth. The magnitude of surge (forward) acceleration during downstrokes was smaller, and that during upstrokes greater, in little penguins than in alcids. This pattern presumably reflected the proportionally greater mass of upstroke muscles in penguins compared with alcids and may allow little penguins to swim at less variable instantaneous speeds.

Animals↗

Body density affects stroke patterns in Baikal seals.

Buoyancy is one of the primary external forces acting on air-breathing divers and it can affect their swimming energetics. Because the body composition of marine mammals (i.e. the relative amounts of lower-density lipid and higher-density lean tissue) varies individually and seasonally, their buoyancy also fluctuates widely, and individuals would be expected to adjust their stroke patterns during dives accordingly. To test this prediction, we attached acceleration data loggers to four free-ranging Baikal seals Phoca sibirica in Lake Baikal and monitored flipper stroking activity as well as swimming speed, depth and inclination of the body axis (pitch). In addition to the logger, one seal (Individual 4) was equipped with a lead weight that was jettisoned after a predetermined time period so that we had a set of observations on the same individual with different body densities. These four data sets revealed the general diving patterns of Baikal seals and also provided direct insights into the influence of buoyancy on these patterns. Seals repeatedly performed dives of a mean duration of 7.0 min (max. 15.4 min), interrupted by a mean surface duration of 1.2 min. Dive depths were 66 m on average, but varied substantially, with a maximum depth of 324 m. The seals showed different stroke patterns among individuals; some seals stroked at lower rates during descent than ascent, while the others had higher stroke rates during descent than ascent. When the lead weight was detached from Individual 4, the seal increased its stroke rate in descent by shifting swimming mode from prolonged glides to more stroke-and-glide swimming, and decreased its stroke rate in ascent by shifting from continuous stroking to stroke-and-glide swimming. We conclude that seals adopt different stroke patterns according to their individual buoyancies. We also demonstrate that the terminal speed reached by Individual 4 during prolonged glide in descent depended on its total buoyancy and pitch, with higher speeds reached in the weighted condition and at steeper pitch. A simple physical model allowed us to estimate the body density of the seal from the speed and pitch (1,027-1,046 kg m(-3), roughly corresponding to 32-41% lipid content, for the weighted condition; 1,014-1,022 kg m(-3), 43-47% lipid content, for the unweighted condition).

Acceleration↗

Regulation of stroke and glide in a foot-propelled avian diver.

Bottom-feeding, breath-hold divers would be expected to minimize transit time between the surface and foraging depth, thus maximizing the opportunities for prey capture during the bottom phase of the dive. To achieve this they can potentially adjust a variety of dive parameters, including dive angle and swim speed. However, because of predictable changes in buoyancy with depth, individuals would also be expected to adjust dive behavior according to dive depth. To test these predictions we deployed miniature, dorsally attached data-loggers that recorded surge and heave accelerations at 64 Hz to obtain the first detailed measurements of a foot-propelled diving bird, the European shag Phalacrocorax aristotelis, in the wild. The results were used to investigate biomechanical changes during the descent, ascent and bottom phases for dives varying between 7 m and 43 m deep. Shags descended and ascended almost vertically (60-90 degrees relative to the sea surface). During descent, swim speed varied between 1.2-1.8 m s(-1) and the frequency of the foot stroke used for propulsion decreased significantly with depth, mainly due to a fivefold increase in the duration of the glide between strokes. Birds appeared to maintain the duration and the maximum strength of power stroke and thus optimize muscle contraction efficiency.

Acceleration↗

Emperor penguins adjust swim speed according to the above-water height of ice holes through which they exit.

Emperor penguins leap from the water onto the sea ice. Their ability to reach above-water height depends critically on initial vertical speed of their leaping, assuming that the kinetic energy is converted to gravitational potential energy. We deliberately changed the above-water heights of ice hole exits, in order to examine whether penguins adjusted swim speed in accordance with the above-water height of the ice. Penguins were maintained in a corral on the fast ice in Antarctica, and voluntarily dived through two artificial ice holes. Data loggers were deployed on the penguins to monitor under water behavior. Nine instrumented penguins performed 386 leaps from the holes during experiments. The maximum swim speeds within 1 s before the exits through the holes correlated significantly with the above-water height of the holes. Penguins adopted higher speed to exit through the higher holes than through the lower holes. Speeds of some failed exits were lower than the theoretical minimum values to reach a given height. Penguins failed to exit onto the sea ice in a total of 37 of the trials. There was no preference to use lower holes after they failed to exit through the higher holes. Rather, swim speed was increased for subsequent attempts after failed leaps. These data demonstrated that penguins apparently recognized the above-water height of holes and adopted speeds greater than the minimal vertical speeds to reach the exit height. It is likely, especially in the case of higher holes (>40 cm), that they chose minimum speeds to exit through the holes to avoid excess energy for swimming before leaping. However, some exceptionally high speeds were recorded when they directly exited onto the ice from lower depths. In those cases, birds could increase swim speed without strokes for the final seconds before exit and they only increased the steepness of their body angles as they surfaced, which indicates that the speed required for leaps by emperor penguins were aided by buoyancy, and that penguins can sometimes exit through the ice holes without any stroking effort before leaping.

Animals↗

Why do macaroni penguins choose shallow body angles that result in longer descent and ascent durations?

It is generally assumed that air-breathing aquatic animals always choose the shortest route to minimize duration for transit between the surface and foraging depth in order to maximize the proportion of time spent foraging. However, empirical data indicate that the body angles of some diving animals are rarely vertical during descent and ascent. Why do they choose shallower body angles that result in longer descent and ascent durations? To investigate this question, we attached acceleration data loggers to eight female macaroni penguins, breeding on the Kerguelen Islands (48 degrees 45'-50 degrees 00' S, 68 degrees 45'-70 degrees 58' E; South Indian Ocean), to record depth, two-dimensional acceleration (stroke cycle frequency and body angle) and temperature. We investigated how they controlled body angle and allocated their submerged time. The instrumented females performed multiple dives (N=6952) with a mean dive depth for each bird ranging from 24.5+/-28.5 m to 56.4+/-75.1 m. Mean body angles during descent and ascent were not vertical. There was large variation in mean descent and ascent angles for a given dive depth, which, in turn, caused large variation in descent and ascent duration. Body angles were significantly correlated with time spent at the bottom-phase of the dive. Birds that spent long periods at the bottom exhibited steep body angles during ascent and subsequent descent. By contrast, they adopted shallow body angles after they had short or no bottom phases. Our results suggest that macaroni penguins stay at the bottom longer after encountering a good prey patch and then travel to the surface at steep body angles. If they do not encounter prey, they discontinue the dive, without staying at the bottom, ascend at shallow body angles and descend at shallow body angles in a subsequent dive. A shallow body angle can increase the horizontal distance covered during a dive, contributing to the move into a more profitable area in the following dive. During the ascent, in particular, macaroni penguins stopped beating their flippers. The buoyantly gliding penguins can move horizontally with minimum stroking effort before reaching the surface.

Acceleration↗

Stroke and glide of wing-propelled divers: deep diving seabirds adjust surge frequency to buoyancy change with depth.

In order to increase locomotor efficiency, breath-holding divers are expected to adjust their forward thrusts in relation to changes of buoyancy with depth. Wing propulsion during deep diving by Brünnich's guillemots (Uria lomvia) was measured in the wild by high-speed (32 Hz) sampling of surge (tail-to-head) and heave (ventral-to-dorsal) accelerations with bird-borne data loggers. At the start of descent, the birds produced frequent surges (3.2 Hz) during both the upstroke and the downstroke against buoyancy to attain a mean speed of 1.2-1.8 m s(-1) that was close to the expected optimal swim speed. As they descended deeper, the birds decreased the frequency of surges to 2.4 Hz, relaying only on the downstroke. During their ascent, they stopped stroking at 18 m depth, after which the swim speed increased to 2.3 m s(-1), possibly because of increasing buoyancy as air volumes expanded. This smooth change of surge frequency was achieved while maintaining a constant stroke duration (0.4-0.5 s), presumably allowing efficient muscle contraction.

Animals↗

Factors affecting stroking patterns and body angle in diving Weddell seals under natural conditions.

Aquatic animals use a variety of strategies to reduce the energetic cost of locomotion. Efficient locomotion is particularly important for breath-holding divers because high levels of exercise may quickly deplete oxygen reserves, leading to the termination of a dive. We investigated the swimming behavior of eight adult Weddell seals, which are proficient divers, in McMurdo Sound, Antarctica. A newly developed data logger was attached to free-ranging females at their own breeding sites to record swimming speed, depth, two-dimensional accelerations (stroke frequency and body angle) and temperature. All seals conducted multiple deep dives (the mean dive depth range for each animal was 223.3+/-66.5-297.9+/-164.7 m). Prolonged gliding while descending was observed with thinner females (N=5 seals). But the fatter females (N=3 seals) exhibited only swim-and-glide swimming, characterized by intermittent stroking and fluctuating swim speed, throughout their descent and ascent. The body angles of four of the seals were restricted to less than 30 degrees by the location of breathing holes in the ice and the slope of local bathymetric features. Of these four, the three fatter seals adopted the stroke-and-glide method while the other thinner seal descended with prolonged periods of gliding. Prolonged gliding seems to be a more efficient method for locomotion because the surface time between dives of gliding seals was significantly less than that of stroking animals, despite their same stroke frequencies.

Acceleration↗

Buoyancy and maximal diving depth in penguins: do they control inhaling air volume?

Using a newly developed data logger to measure acceleration, we demonstrate that free-ranging king and Adélie penguins only beat their flippers substantially during the first part of descent or when they were presumed to be chasing prey at the bottom of dives. Flipper beating stopped during the latter part of ascent: at 29+/-9 % (mean +/- S.D.) of dive depth (mean dive depth=136.8+/-145.1 m, N=425 dives) in king penguins, and at 52+/-20 % of dive depth (mean dive depth=72.9+/-70.5 m, N=664 dives) in Adélie penguins. Propulsive swim speeds of both species were approximately 2 m s(-1) during dives; however, a marked increase in speed, up to approximately 2.9 m s(-1), sometimes occurred in king penguins during the passive ascending periods. During the prolonged ascending, oblique ascent angle and slowdown near the surface may represent one way to avoid the potential risk of decompression sickness. Biomechanical calculations for data from free-ranging king and Adélie penguins indicate that the air volume of the birds (respiratory system and plumage) can provide enough buoyancy for the passive ascent. When comparing the passive ascents for shallow and deep dives, there is a positive correlation between air volume and the depth of the dive. This suggests that penguins regulate their air volume to optimize the costs and benefits of buoyancy.

Air↗