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1.
The energetic costs of swimming at the surface (swimming) and swimming underwater (diving) are compared in tufted ducks (Aythya fuligula) and three species of penguins, the gentoo (Pygoscelis papua), the king (Aptenodytes patagonicus), and the emperor (Aythya forsteri). Ducks swim on the surface and use their webbed feet as paddles, whereas penguins tend to swim just below the surface and use their flippers as hydrofoils, the latter being much more efficient. Penguins are more streamlined in shape. Thus, the amount of energy required to transport a given mass of bird a given distance (known as the cost of transport) is some two to three times greater in ducks than in penguins. Ducks are also very buoyant, and overcoming the force of buoyancy accounts for 60% and 85% of the cost of descent and remaining on the bottom, respectively, in these birds. The energy cost of a tufted duck diving to about 1.7 m is similar to that when it is swimming at its maximum sustainable speed at the surface (i.e., approximately 3.5 times the value when resting on water). Nonetheless, because of the relatively short duration of its dives, the tufted duck dives well within its calculated aerobic dive limit (cADL, usable O(2) stores per rate of O(2) usage when underwater). However, these three species of penguins have maximum dive durations ranging from 5 min to almost 16 min and maximum dive depths from 155 to 530 m. When these birds dive, they have to metabolise at no more than when resting in water in order for cADL to encompass the duration of most of their natural dives. In gentoo and king penguins, there is a fall in abdominal temperature during bouts of diving; this may reduce the oxygen requirements in the abdominal region, thus enabling dive duration to be extended further than would otherwise be the case.  相似文献   

2.
To better understand how elephant seals (Mirounga angustirostris) use negative buoyancy to reduce energy metabolism and prolong dive duration, we modelled the energetic cost of transit and deep foraging dives in an elephant seal. A numerical integration technique was used to model the effects of swim speed, descent and ascent angles, and modes of locomotion (i.e. stroking and gliding) on diving metabolic rate, aerobic dive limit, vertical displacement (maximum dive depth) and horizontal displacement (maximum horizontal distance along a straight line between the beginning and end locations of the dive) for aerobic transit and foraging dives. Realistic values of the various parameters were taken from previous experimental data. Our results indicate that there is little energetic advantage to transit dives with gliding descent compared with horizontal swimming beneath the surface. Other factors such as feeding and predator avoidance may favour diving to depth during migration. Gliding descent showed variable energy savings for foraging dives. Deep mid-water foraging dives showed the greatest energy savings (approx. 18%) as a result of gliding during descent. In contrast, flat-bottom foraging dives with horizontal swimming at a depth of 400m showed less of an energetic advantage with gliding descent, primarily because more of the dive involved stroking. Additional data are needed before the advantages of gliding descent can be fully understood for male and female elephant seals of different age and body composition. This type of data will require animal-borne instruments that can record the behaviour, three-dimensional movements and locomotory performance of free-ranging animals at depth.  相似文献   

3.

Background

Because they have air stored in many body compartments, diving seabirds are expected to exhibit efficient behavioural strategies for reducing costs related to buoyancy control. We study the underwater locomotor activity of a deep-diving species from the Cormorant family (Kerguelen shag) and report locomotor adjustments to the change of buoyancy with depth.

Methodology/Principal Findings

Using accelerometers, we show that during both the descent and ascent phases of dives, shags modelled their acceleration and stroking activity on the natural variation of buoyancy with depth. For example, during the descent phase, birds increased swim speed with depth. But in parallel, and with a decay constant similar to the one in the equation explaining the decrease of buoyancy with depth, they decreased foot-stroke frequency exponentially, a behaviour that enables birds to reduce oxygen consumption. During ascent, birds also reduced locomotor cost by ascending passively. We considered the depth at which they started gliding as a proxy to their depth of neutral buoyancy. This depth increased with maximum dive depth. As an explanation for this, we propose that shags adjust their buoyancy to depth by varying the amount of respiratory air they dive with.

Conclusions/Significance

Calculations based on known values of stored body oxygen volumes and on deep-diving metabolic rates in avian divers suggest that the variations of volume of respiratory oxygen associated with a respiration mediated buoyancy control only influence aerobic dive duration moderately. Therefore, we propose that an advantage in cormorants - as in other families of diving seabirds - of respiratory air volume adjustment upon diving could be related less to increasing time of submergence, through an increased volume of body oxygen stores, than to reducing the locomotor costs of buoyancy control.  相似文献   

4.
5.
Synchronizing behavior with other conspecifics has been suggested as serving a function of increased foraging efficiency. However, the potential costs associated with synchronization of behavior have rarely been studied. Adélie penguins Pygoscelis adeliae sometimes dive synchronously in small open waters surrounded by fast sea ice. We examined the diving behavior of three couples and one trio, which were observed to dive synchronously among groups of 12–47 birds for 1.7–4.5 h duration, with time-depth recorders. Timing of diving and surfacing differed slightly between individuals, and one bird tended to initiate diving earlier than the other. Although the duration of the dives differed only slightly between these birds, the maximum depth of the dives differed to a large extent, with one member tending to dive consistently deeper than the other bird in two out of the four cases. Vertical distances between tagged birds in the undulatory phases of the dives (presumed feeding time) were greater than those in the descent and ascent phases, suggesting independent foraging by group members. Duration of the undulatory phase of the dives tended to be shorter in deeper-diving individuals than the others in the synchronously diving group, suggesting a potential cost of reduced feeding time to synchronize diving and surfacing with other birds. A digital video image relating to the article is available at .  相似文献   

6.
We attached a video system and data recorder to a northern elephant seal to track its three-dimensional movements and observe propulsive strokes of the hind flippers. During 6 h of recording, the seal made 20 dives and spent 90% of the time submerged. Average dive duration, maximum depth and swimming speed were 14.9 min+/-6.1 S.D., 289 m+/-117 S.D. and 1.1 m s(-1)+/-0.12 S.D., respectively. The distance swum during a dive averaged 925 m+/-339 S.D., and the average descent and ascent angles were 41 degrees +/-18 S.D. and 50 degrees +/-21 S.D., respectively. Dive paths were remarkably straight suggesting that the seal was navigating while submerged. We identified three modes of swimming based on the interval between propulsive strokes: continuous stroking; stroke-and-glide swimming; and prolonged gliding. The seal used continuous stroking from the surface to a mean depth of 20 m followed by stroke-and-glide swimming. Prolonged gliding started at a mean depth of 60 m and continued to the bottom of dives. For dives to depths of 300 m or more, 75% of the descent time was spent in prolonged gliding and 10% in stroke-and-glide swimming, amounting to 5.9-9.6 min of passive descent per dive. Average swimming speed varied little with swimming mode and was not a good indicator of propulsive effort. It appears that the seal can use prolonged gliding to reduce the cost of transport and increase dive duration. Energetically efficient locomotion may help explain the long and deep dives that routinely exceed the theoretical aerobic dive limit in this species.  相似文献   

7.
Semi‐aquatic mammals have secondarily returned to the aquatic environment, although they spend a major part of their life operating in air. Moving both on land, as well as in, and under water is challenging because such species are considered to be imperfectly adapted to both environments. We deployed accelerometers combined with a depth sensor to study the diving behavior of 12 free‐living Eurasian beavers Castor fiber in southeast Norway between 2009 and 2011 to examine the extent to which beavers conformed with mass‐dependent dive capacities, expecting them to be poorer than wholly aquatic species. Dives were generally shallow (<1 m) and of short duration (<30 s), suggesting that the majority of dives were aerobic. Dive parameters such as maximum diving depth, dive duration, and bottom phase duration were related to the effort during different dive phases and the maximum depth reached. During the descent, mean vectorial dynamic body acceleration (VeDBA—a proxy for movement power) was highest near the surface, probably due to increased upthrust linked to fur‐ and lung‐associated air. Inconsistently though, mean VeDBA underwater was highest during the ascent when this air would be expected to help drive the animals back to the surface. Higher movement costs during ascents may arise from transporting materials up, the air bubbling out of the fur, and/or the animals’ exhaling during the bottom phase of the dive. In a manner similar to other homeotherms, beavers extended both dive and bottom phase durations with diving depth. Deeper dives tended to have a longer bottom phase, although its duration was shortened with increased VeDBA during the bottom phase. Water temperature did not affect diving behavior. Overall, the beavers’ dive profile (depth, duration) was similar to other semi‐aquatic freshwater divers. However, beavers dived for only 2.8% of their active time, presumably because they do not rely on diving for food acquisition.  相似文献   

8.
Summary Humboldt Penguins diving in clear water in Chile had two distinct behaviours; short dives where the birds remained within a metre of the surface and bounce dives, where the penguins descended the water column to the sea bed where they immediately returned to the surface. Here, dive time was correlated with water depth. Stomach pumped penguins had eaten predominantly pelagic school fish,Scomberesox, Engraulis andSardinops. Most prey (94 %) had been seized from below. We suggest that short, shallow dives are typical of travelling penguins. Bounce dives, however, would enable foraging penguins, which have an enhanced binocular field above the line of the bill, to perceive surface-swimming prey more easily because silvery clupeid fish appear as silhouettes.
Zusammenfassung In klaren Gewässern in Chile tauchende Humboldt-Pinguine zeigten zwei deutlich verschiedene Verhaltensweisen. Während kurzer Tauchgänge blieben die Tiere im Bereich von 1 m unter der Oberfläche. Tauchgänge durch unmittelbares Abtauchen in einem Winkel von 45° eingeleitet, führten direkt zum Meeresboden; die Vögel kehrten sofort wieder an die Oberfläche zurück. In diesem Fall war die Tauchzeit von der Wassertiefe abhängig. Pinguine, deren Magen durch Auspumpen untersucht wurde, hatten vorwiegend pelagische Schwarmfische gefressen:Scomberesox, Engraulis undSardinops. Die Mehrzahl der Fische (94 %) wurde von unten kommend erfaßt. Nach unserer Auffassung sind die kurzen und flachen Tauchgänge typisch für nicht jagende Pinguine, die längere horizontale Strecken zurücklegen. Das tiefe Abtauchen dagegen ermöglicht jagenden Pinguinen, die ein besonders gutes Fernsichtvermögen oberhalb des Schnabels besitzen, nahe der Oberfläche schwimmende Beute leichter zu erfassen. Die sonst silbrigen clupeiden Fische sind als dunkle Silhouetten gegen die Wasseroberfläche besser zu erkennen.
  相似文献   

9.
Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water<5 m deep at two sites. Dive duration increases with water depth at both sites, but the relationship is weak. Dives were longer than dives by cormorants in water of similar depth (max 108 s in water 2.5 m deep), with dives of up to 68 s observed in water<0.5 m deep. Initial dives in a bout were shorter than expected, possibly because their plumage was not fully saturated. Dive efficiency (dive:rest ratio) was 5–6, greater than cormorants (2.7±0.4 for 18 species) and other families of diving birds (average 0.2–4.3). Post-dive recovery periods increased with dive duration, but only slowly, resulting in a strong increase in efficiency with dive duration. All dives are likely to fall within the theoretical anaerobic dive limit. Foraging bouts were short (17.8±4.3 min) compared to cormorants, with birds spending 80±5% of time underwater. Darters take advantage of their low buoyancy to forage efficiently in shallow water, and their slow, stealthy dives are qualitatively different from those of other diving birds. However, they are forced to limit the duration of foraging bouts by increased thermoregulatory costs associated with wettable plumage.  相似文献   

10.
We collected simultaneous dive Time Depth Recorder (TDR) data and video images from free swimming adult female leatherback turtles, Dermochelys coriacea, during the first 24 h after nesting on the beach, in order to determine relationships between dive parameters, activity, overall respiratory frequency and behaviour.We identified three different underwater locomotory activities (subsurface swimming, V-shaped dives and U-shaped dives) from video and TDR data that varied in their mean depth, duration and a number of other parameters. Overall respiratory frequency (overall fR) was significantly different between all locomotory activities, with turtles taking 1.7±0.1 breaths min−1 while subsurface swimming, 0.78 breaths min−1 after V-shaped dives and 0.57 breaths min−1 after U-shaped dives. Descent rates and ascent rates were significantly faster in U-shaped dives (descent 0.19±0.010 m s−1, ascent 0.28±0.015 m s−1) than in V-shaped dives (descent 0.10±0.008 m s−1, ascent 0.12±0.012 m s−1). Flipper stroke rates were significantly lower during the bottom component of U-shaped dives (0.18±0.02 strokes s−1) than during the descent (0.29±0.03 strokes s−1) or ascent (0.29±0.03 strokes s−1). From overall fR and flipper stroke rate data, we inferred that turtles used less energy during U-shaped dives than the other activity types. We recorded interactions between male turtles and the study females that lasted up to 11 min, during which male turtles displayed the characteristic courtship behaviour of sea turtles. It appeared that females attempted to avoid males by aborting ascent and extending dive duration to swim to the sea floor when males were present.  相似文献   

11.
Cormorants hunt both benthic (sedentary) and pelagic (motile) prey but it is not known if the energy costs of foraging on these prey differ. We used respirometry to measure the costs of diving in double-crested cormorants (Phalacrocorax auritus) foraging either for sedentary (fish pieces) or motile (juvenile salmon) prey in a deep dive tank. Short dives for sedentary prey were more expensive than dives of similar duration for motile prey (e.g. 20% higher for a 10s dive) whereas the reverse was true for long dives (i.e. long dives for motile prey were more expensive than for sedentary prey). Across dives of all durations, the foraging phase of the dive was more expensive when the birds hunted motile prey, presumably due to pursuit costs. The period of descent in all the dives undertaken appears to have been more expensive when the birds foraged on sedentary prey, probably due to a higher swimming speed during this period.  相似文献   

12.
The diving behaviour of the Shy Albatross Diomedea cauta was investigated using archival time-depth recorders (TDRs) and maximum depth gauges (MDGs). Data from birds carrying multiple devices and from diving simulations indicated that the degree of correspondence between TDRs and MDGs varied with the dive depth, duration and frequency, as well as with body placement. The MDGs were the most reliable when the diving depth was greater than 0.5 m, when the diving frequency was low and when gauges were placed on the birds' backs. The TDRs were used during late incubation and early chick rearing in 1994. Fifty-two dives (0.4 m) were recorded during 20 foraging trips of 15 individuals. The majority of dives were within the upper 3 m of the water column and lasted for less than 6 s. However, dives to 7.4 m and others lasting 19 s were recorded. The albatrosses dived between 07.00 h and 22.00 h, with peaks in their diving activity near midday and twilight. Mean diving depth varied throughout the day. with the deepest dives occurring between 10.00 h and 12.00 h. Two dive types were identified on the basis of the relationship between dive depth and descent rate. Plunge dives were short (5 s), and the birds reached a maximum depth of 2.9 m. Swimming dives were both longer and deeper. The characteristics of Shy Albatross plunge dives were similar to those of gannets Morus spp., which are known to be proficient plunge divers. Swimming dives suggest that Shy Albatrosses actively pursue prey underwater.  相似文献   

13.
In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.  相似文献   

14.
Data on the swim speed, dive depth and feeding rates of three Adélie penguins (Pygoscelis adeliae) foraging in summer 1998/1999 in Adélie Land, Antarctica were collected using dorsally-mounted loggers, in tandem with oesophageal temperature sensors. Swim speed could be integrated, together with the rate of change of depth, to determine dive and return-to-surface angles. Overall, birds increased rates of change of depth during commuting phases so that dive angles were steeper in dives terminating at greater depths. Angles of descent and ascent during feeding dives were greater than during non-feeding dives. Variation in the descent angle over time of particular dives was generally less than 10°, but the angles of the ascent phases varied more widely. The importance of selecting the optimum descent and ascent angles with respect to prey exploitation, oxygen stores and time gained in the feeding area over the course of a dive by diving at a steeper angle is discussed.  相似文献   

15.
Physical environment and physiological characteristics of marine mammals potentially affect the duration and depth of diving. Härkönen (1987b) proposed a hypothesis that the harbor seal would gain maximum energy by foraging at intermediate depths. To investigate this hypothesis, we studied diving behavior of the Pacific harbor seal (Phoca vitulina ricbardii) during 1995 through 1997 in Monterey Bay, California. Dive depths (n = 13,063 dives) were recorded via time‐depth recorders. Approximately 80% of recorded dives were classified as square dives (type I), which typically were associated with foraging in pinnipeds. Approximately 11% of dives were V dives (type II; 1,402 dives), and the remainder (1,225 dives) were skewed dives (type III and IV). The deepest recorded dive was 481 m, while the greatest duration was 35.25 min. Body mass explained the variability of durations of long dives for females (95th percentile; D95♂=‐5.47 + 0.18 × (mass♀), r2= 0.91, 95% CI for slope = [0.08, 0.28], n= 5) and for males (D95♂=‐5.86 + 0.18 × (mass♀), r2= 0.83, 95% CI for slope = [0.12, 0.24], n= 11). The large proportion of variability in deep dives, however, was explained by body mass only for males (95th percentile; Z95♂=‐363.9 + 6.05 × (mass♀), r2= 0.83, 95% CI for slope = [3.93, 8.17], n= 11) and not for females (Z95,♂=?148.1 +3.11 × (mass♀), r2= 0.58, 95% CI for slope = [‐1.7, 7.9], n= 5, 95% CI for slope= [?1.7, 7.9]). Median depths of presumed foraging dives of harbor seals in the Monterey Bay area were between 5 and 100 m, which were within the range of the previously reported depths for other areas (< 100 m). Our findings generally supported Härkönen's hypothesis that harbor seals forage in the intermediate depth in their environment.  相似文献   

16.
Three juvenile narwhals captured during August 1998 in the northeast of Svalbard, Norway, were equipped with satellite-relayed data loggers (SRDLs) that transmitted diving and swim-speed data, in addition to location, for up to 46 days. A total of 1,354 complete dive cycles were recorded. Most of the diving was shallow and of short duration. Maximum recorded dive depth was 546 m, maximum recorded dive duration was 24.8 min, and maximum recorded swim-speed was 4.7 ms−1. Ascent speed, vertical ascent speed, descent speed and vertical descent speed were all significantly higher during deep dives (>200 m) than for shallow dives (<200 m). In addition both ascent and descent angles were much steeper for deep dives than during shallow dives. Most of the shallow diving seemed to be associated with travelling, with the animal shifting between various locations, while the deep diving (often to the bottom) for extended periods in some specific areas might have been associated with foraging. Even though the sample size in this study is small, the data are the first information available for movements and diving behaviour of narwhals near Svalbard.  相似文献   

17.
J. P. Croxall    Y. Naito    A. Kato    P. Rothery    D. R. Briggs 《Journal of Zoology》1991,225(2):177-199
The pattern and characteristics of diving of two male blue-eyed shags Phalacrocorax atriceps were studied, using continuous-recording time-depth recorders, for a total of 15 consecutive days during which the depth, duration, bottom time, ascent and descent rates and surface intervals of 674 dives were recorded. Deep dives (> 35 m, averages80–90 m, max. 116 m) were twice as common (64% versus 34%) as shallow dives (< 21 m and 90% < 10 m). Deep dives were long (averages 2.7-4.1 min, max. 5.2 min) with half the time spent near maximum depth and fast travel speeds (averages 1.0-2.4 m s−1). Shallow dives were short (average 0.5 min, max. 1.3 min), without bottom time and with slow travel speeds (0.1–0.6 m s−1). The time spent at depth and the diet (mainly benthic fish and octopus) is consistent with benthic foraging; the function of shallow dives is uncertain. Male shags forage mainly in the afternoon in3–5 distinct bouts of diving. Within bouts (and shorter homogeneous sequences of diving) surface intervals are consistently2–3 times the preceding dive duration; in other shags the reverse is the case. Blue-eyed shag diving depth, duration and pattern is extreme amongst shags; and the relationship between dives and surface intervals suggests that they may regularly exceed their aerobic dive limit.  相似文献   

18.
The underwater behaviour of 11 belugas or white whales was examined during summer using time-at-depth records relayed by satellite-linked data-loggers. Simultaneous tracking information was obtained for each whale. Eight distinct dive profiles were identified in submergences made to depths of >40 m. Four of these, together comprising 84% of these “deep” dives, were of a square profile. They were characterised by a continuous descent to a particular depth (usually the sea bed), a “bottom phase” at or near that depth, and a direct ascent to the surface. These dives are presumed to be made for benthic foraging. Other, much less common, dive shapes were “V”-shaped, parabolic and trapezoidal. “Shallow” dives (15–40 m depth) were of a variety of shapes, short duration and high average horizontal speeds. Many probably occurred during periods of directed travel. This population of belugas treats most of the water column as dead space separating resources of oxygen and nutrition. Received: 8 January 1998 / Accepted: 27 April 1998  相似文献   

19.
We investigated the diving behaviour, the time allocation of the dive cycle and the behavioural aerobic dive limit (ADL) of platypuses (Ornithorhynchus anatinus) living at a sub-alpine Tasmanian lake. Individual platypuses were equipped with combined data logger-transmitter packages measuring dive depth. Mean dive duration was 31.3 s with 72% of all dives lasting between 18 and 40 s. Mean surface duration was 10.1 s. Mean dive depth was 1.28 m with a maximum of 8.77 m. Platypuses performed up to 1600 dives per foraging trip with a mean of 75 dives per hour. ADL was estimated by consideration of post-dive surface intervals vs. dive durations. Only 15% of all dives were found to exceed the estimated ADL of 40 s, indicating mainly aerobic diving in the species. Foraging platypuses followed a model of optimised recovery time, the optimal breathing theory. Total bottom duration or total foraging duration per day is proposed as a useful indicator of foraging efficiency and hence habitat quality in the species.  相似文献   

20.
To be successful, marine predators must alter their foraging behavior in response to changes in their environment. To understand the impact and severity of environmental change on a population it is necessary to first describe typical foraging patterns and identify the underlying variability that exists in foraging behavior. Therefore, we characterized the at‐sea behavior of adult female California sea lions (n = 32) over three years (2003, 2004, and 2005) using satellite transmitters and time‐depth recorders and examined how foraging behavior varied among years. In all years, sea lions traveled on average 84.7 ± 11.1 km from the rookery during foraging trips that were 3.2 ± 0.3 d. Sea lions spent 42.7% ± 1.9% of their time at sea diving and displayed short (2.2 ± 0.2 min), shallow dives (58.5 ± 8.5 m). Among individuals, there was significant variation in both dive behavior and movement patterns, which was found in all years. Among years, differences were found in trip durations, distances traveled, and some dive variables (e.g., dive duration and bottom time) as sea lions faced moderate variability in their foraging habitat (increased sea‐surface temperatures, decreased upwelling, and potential decreased prey abundance). The flexibility we found in the foraging behavior of California sea lions may be a mechanism to cope with environmental variability among years and could be linked to the continuing growth of sea lion populations.  相似文献   

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