Metabolic rates and the energetic cost of external tag attachment in juvenile blacktip sharks Carcharhinus limbatus

This study reports on the metabolic rate of the blacktip shark Carcharhinus limbatus and the energetic costs of external tag attachment. Metabolic rates, swimming speed and tail‐beat (B_T) frequency were measured in a static respirometer with untagged animals and animals equipped with a small data logger. Tagged sharks showed significantly higher routine oxygen consumption and lower swimming speeds than untagged animals, indicating that tagging significantly affected the swimming efficiency and energetic requirements in these small sharks, and that these effects must be accounted for when interpreting telemetry data from free‐ranging individuals.     

Rainbow trout <Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis> energetic responsesto pulsed flows in the American River,California, assessed by electromyogram telemetry

Although rainbow trout Oncorhynchus mykiss within the American River, California, apparently exhibit minimal upstream or downstream movements in response to hydroelectric-power-generation-related pulsed flows, the associated energetic costs are unknown. We implanted rainbow trout (n = 9, ≥30 cm SL) with electromyogram (EMG)-sensor-equipped radio transmitters to assess the swimming behavior and associated energetic costs associated with their responses to pulsed flows. Using laboratory calibrations in a Brett-type swimming respirometer, the trouts’ swimming speeds and oxygen consumption rates were estimated for their in-river EMG data, through a complete hydroelectric power-generation river pulsed-flow sequence (pre-pulse, increasing flow, peak, and decreasing flow stages), on several (mean: 3.2) sampling dates. Using a mixed-linear model, we found that fish swimming speed estimates increased during the increasing flow stage, while the associated mean oxygen consumption rates also increased at this stage. At river flows near the usual peak (>44 m³s⁻¹), swimming speeds and movement rates decreased, possibly due to the fish using the river’s habitat complexities as hydraulic cover. We conclude that rainbow trout incur increased swimming-related energetic costs during increasing flows and, potentially, decreased foraging opportunities at high flows.     

The influence of depth on a breath-hold diver: Predicting the diving metabolism of Steller sea lions (Eumetopias jubatus)

Diving animals must endeavor to increase their dive depths and prolong the time they spend exploiting resources at depth. Results from captive and wild studies suggest that many diving animals extend their foraging bouts by decreasing their metabolisms while submerged. We measured metabolic rates of Steller sea lions (Eumetopias jubatus) trained to dive to depth in the open ocean to investigate the relationships between diving behaviour and the energetic costs of diving. We also constructed a general linear model to predict the oxygen consumption of sea lions diving in the wild. The resultant model suggests that swimming distance and depth of dives significantly influence the oxygen consumption of diving Steller sea lions. The predictive power of the model was tested using a cross-validation approach, whereby models reconstructed using data from pairs of sea lions were found to accurately predict the oxygen consumption of the third diving animal. Predicted oxygen consumption during dives to depth ranged from 3.37 L min^− 1 at 10 m, to 1.40 L min^− 1 at 300 m over a standardized swimming distance of 600 m. This equated to an estimated metabolic rate of 97.54 and 40.52 MJ day^− 1, and an estimated daily feeding requirement of 18.92 and 7.96 kg day^− 1 for dives between 10 and 300 m, respectively. The model thereby provides information on the potential energetic consequences that alterations in foraging strategies due to changes in prey availability could have on wild populations of sea lions.     

Swimming speed, respiration rate, and estimated cost of transport in adult killer whales

The physiology of free-ranging cetaceans is difficult to study and as a consequence, data on the energetics of these animals are limited. To better understand the energetic cost of swimming in killer whales, total cost of transport ( COT ) was estimated from swimming speeds and respiration rates from wild adult northern resident killer whales ( Orcinus orca ) and reported values of oxygen consumption in captive whales. Respiration rate (breaths per minute) was positively correlated with swimming speed (meters per second), while mass-specific COT (Joules per kilogram per meter) decreased with speed. Lack of data on very fast-swimming animals hindered assessment of the exact speed at which COT was minimal. However, minimum mass-specific COT for killer whales in the present study approached those predicted by a previously published allometric equation for marine mammals, and corresponded to "optimal" swimming speeds of 2.6–3 m/s. Interestingly, the observed average swimming speed (1.6 m/s) was lower than predicted optimal swimming speed. Finally, females with dependent calves had higher respiration rates than females without calves. These findings could be due to synchronous breathing with calves or could result from increased costs of lactation and swimming with a calf in echelon formation. Consequently, females with calves may have much greater COT at optimal swimming speeds than females without calves.     

Homing ability of young lemon sharks, <Emphasis Type="Italic">Negaprion brevirostris</Emphasis>

Synopsis We carried out the first experimental study testing an elasmobranchs ability to return home. We displaced juvenile lemon sharks,Negaprion brevirostris, 4–16 km from their observed home ranges at Bimini Islands, Bahamas during daylight and at night. We tracked all sharks except one back to the Bimini Islands and most returned to their home ranges observed before displacement. Even sharks displaced to a site closer to another island with suitable habitat for young lemon sharks returned to their home ranges at Bimini Islands. Sharks displayed a preferred compass direction (PCD) toward the east as their first swimming direction after release, suggesting an innate sense of direction. This bearing was followed shortly afterwards by a home-oriented direction. Swimming speeds prior to reaching shore were approximately twice as fast than the usual cruising speed reported for juvenile lemon sharks. The return of young (0–2 years), inexperienced sharks to their original home range indicate high site fidelity and an ability to home.     

Behavioural basis of intertidal zonation in Eurydice pulchra Leach

Eurydice pulchra Leach freshly collected at neap and spring tides exhibits an endogenous circatidal rhythm of swimming when kept without sand in constant conditions in laboratory aquaria. When provided with sand in the laboratory, tidally-phased swimming may occur spontaneously at some times of the lunar month, predominantly after the times of highest spring tides but not at neaps. In these cases, therefore, there appears to be clearly demonstrated for the first time in a marine animal a circasemilunar pattern of emergence from the substratum which is coupled to a circatidal rhythm of swimming; the isopods in the laboratory emerge from the sediment only when the swimming rhythm and emergence rhythm are at their maxima. In the field at neap tides some isopods emerge to swim at high tide, dependent upon their responses to water agitation, their feeding state and responses to light. At and just after spring tides in the field large numbers of the isopods swim at high tide, triggered by the endogenous circasemilunar rhythm of emergence which in the field is probably cued by the effects of increased water agitation. The main function of the endogenous circatidal rhythm of swimming appears to be to permit the isopods to return to their preferred zone in the sand before the fall of the tide. The coupled circasemilunar rhythm of emergence which induces greater swimming at spring tides reduces the risk of animals being stranded high on the shore during neap tides.     

Tidal movements of female leopard sharks (<Emphasis Type="Italic">Triakis semifasciata</Emphasis>) in Elkhorn Slough,California

The leopard shark (Triakis semifasciata) is one of the most common species of elasmobranch in California, and uses the shallow bays and estuaries of California extensively throughout its life history. To examine the role that tides and time of day play on the distribution and movements of leopard sharks in an estuarine environment, a total of 22 female leopard sharks (78–140 cm TL) were tagged with acoustic transmitters in Elkhorn Slough, California, USA. Eight sharks were manually tracked for 20–71.5 h, and 13 sharks were monitored for 4–280 days using an array of acoustic receivers. Overall, the distribution and movements of sharks were strongly influenced by the tides and to a lesser extent by period of day, although general patterns of movement differed depending on what region of Elkhorn Slough the sharks were using. In the main channel of Elkhorn Slough, sharks generally moved with the tide, maximizing the area over which they could forage on a more dispersed prey field. Conversely, leopard sharks within the Elkhorn Slough National Estuarine Research Reserve regularly swam against strong currents to remain in proximity to the intertidal mudflats. This high degree of fidelity to a specific region was probably due to an abundance of important prey in the area. These results indicate that movements, and thus the foraging ecology, of leopard sharks show a high degree of plasticity and are influenced by tidal stage, tidal current, availability of suitable habitat, and availability and distribution of important prey items.     

The energetic costs of alternative male reproductive strategies in <Emphasis Type="Italic">Xiphophorus nigrensis</Emphasis>

The coexistence of alternative male mating strategies depends on the balance between costs and benefits. Here we examine the short-term metabolic costs associated with distinct reproductive strategies in the genetically determined alternative male phenotypes of a northern swordtail, Xiphophorus nigrensis. In this species, large males court females, non-adorned small males chase females, and intermediate males exhibit both courtship and chase behaviors. Using intermittent flow respirometry, we measure oxygen consumption rates and behaviors of each size class in isolation and in the presence of a female. Changes in oxygen consumption between solitary and female presence trials (ΔVO₂) correlated significantly with standard length across all size classes (r = 0.42). Only the large male class exhibited a significant increase in oxygen consumption in female-present trials exhibiting a range of increase from 2 to 200% relative to solitary metabolic rates, but costs of specific courtship displays could not be demonstrated. Sword length explained 54–57% of the variation in oxygen consumption in large male solitary trials and 63–65% in the female-present trials independent of any behavioral correlation with sword length. Our results exhibit similarities to condition-dependent alternative mating systems where the female-favored phenotype has higher energetic costs.     

Effect of weight and frontal area of external telemetry packages on the kinematics,activity levels and swimming performance of small‐bodied sharks

This study sought to observe the effects of submerged weight and frontal cross‐sectional area of external telemetry packages on the kinematics, activity levels and swimming performance of small‐bodied juvenile sharks, using lemon sharks Negaprion brevirostris (60–80 cm total length, L_T) as a model species. Juveniles were observed free‐swimming in a mesocosm untagged and with small and large external accelerometer packages that increased frontal cross‐sectional area of the animals and their submerged weight. Despite adhering to widely used standards for tag mass, the presence of an external telemetry package altered swimming kinematics, activity levels and swimming performance of juvenile N. brevirostris relative to untagged individuals, suggesting that tag mass is not a suitable standalone metric of device suitability. Changes in swimming performance could not be detected from tail‐beat frequency, which suggests that tail‐beat frequency is an unsuitable standalone metric of swimming performance for small N. brevirostris. Lastly, sharks experienced treatment‐specific changes in activity level and swimming kinematics from morning to afternoon observation. Therefore, the presence of external telemetry packages altered the kinematics, activity levels and swimming performance of small young‐of‐the‐year N. brevirostris and these data may therefore be relevant to other similar‐sized juveniles of other shark species.     

Bioenergetics of free-ranging juvenile scalloped hammerhead sharks (Sphyrna lewini) in Kāne'ohe Bay, ō'ahu, HI

Metabolic and activity rates determined from free-ranging juvenile scalloped hammerhead shark pups (Sphyrna lewini) in Kāne'ohe Bay, ō'ahu, HI, were used to develop a simplified energy budget for this population. Five shark pups were tracked using a specially designed acoustic tail beat transmitter. Previous laboratory experiments determined that tail beat frequency (TBF) and water temperature could be used as predictors of instantaneous swimming speed (U) and oxygen consumption rates (V?o₂). Sharks carrying transmitters had higher cost of transport than uninstrumented sharks, but because the difference was quantifiable, appropriate corrections could be made for transmitter effects on energy consumption of instrumented sharks. Sharks tracked in Kāne'ohe Bay had an overall average TBF of 70±10 beats min⁻¹ and a U of 0.81±0.1 body lengths s⁻¹, but swam significantly faster at night than during the day. These sharks also exhibited high average metabolic rates (MR) (96±15 kJ kg⁻¹ day⁻¹) compared with other species of sharks previously studied. Sharks tracked during warmer summer months swam slightly faster and exhibited higher MR than one tracked in December. The high MR measured for sharks in the bay indicate that these sharks require a high daily ration. The relatively low caloric value of the most common prey (snapping shrimp, Alpheus malabaricus) suggests that high numbers must be consumed to meet the sharks' daily requirements. Low and/or negative growth rates of shark pups in the field and declining population size over the summer season suggests that a significant percentage of pups in Kāne'ohe Bay may starve as the result of their high MR requirements. Although availability of A. malabaricus may not be limiting in Kāne'ohe Bay, foraging skills of some neonates may be inadequate to enable them to meet their daily energetic needs. However, sharks that are successful in surviving through the winter may actually grow faster in the colder months due to the temperature dependent decrease of their MR and a reduction in conspecific competition as the result of high summer seasonal attrition due to starvation and emigration.     

Unexpected Positive Buoyancy in Deep Sea Sharks,Hexanchus griseus,and a Echinorhinus cookei

We do not expect non air-breathing aquatic animals to exhibit positive buoyancy. Sharks, for example, rely on oil-filled livers instead of gas-filled swim bladders to increase their buoyancy, but are nonetheless ubiquitously regarded as either negatively or neutrally buoyant. Deep-sea sharks have particularly large, oil-filled livers, and are believed to be neutrally buoyant in their natural habitat, but this has never been confirmed. To empirically determine the buoyancy status of two species of deep-sea sharks (bluntnose sixgill sharks, Hexanchus griseus, and a prickly shark, Echinorhinus cookei) in their natural habitat, we used accelerometer-magnetometer data loggers to measure their swimming performance. Both species of deep-sea sharks showed similar diel vertical migrations: they swam at depths of 200–300 m at night and deeper than 500 m during the day. Ambient water temperature was around 15°C at 200–300 m but below 7°C at depths greater than 500 m. During vertical movements, all deep-sea sharks showed higher swimming efforts during descent than ascent to maintain a given swimming speed, and were able to glide uphill for extended periods (several minutes), indicating that these deep-sea sharks are in fact positively buoyant in their natural habitats. This positive buoyancy may adaptive for stealthy hunting (i.e. upward gliding to surprise prey from underneath) or may facilitate evening upward migrations when muscle temperatures are coolest, and swimming most sluggish, after spending the day in deep, cold water. Positive buoyancy could potentially be widespread in fish conducting daily vertical migration in deep-sea habitats.     

Review of Elasmobranch Behavioral Studies Using Ultrasonic Telemetry with Special Reference to the Lemon Shark, Negaprion Brevirostris, Around Bimini Islands, Bahamas

A review of past behavioral ultrasonic telemetry studies of sharks and rays is presented together with previously unpublished material on the behavior of the lemon shark, Negaprion brevirostris, around the Bimini Islands, Bahamas. The review, focusing on movement behaviors of 20 shark and three ray species, reveals that elasmobranchs exhibit a variety of temporal and spatial patterns in terms of rates-of-movement and vertical as well as horizontal migrations. The lack of an apparent pattern in a few species is probably attributable to the scarcity of tracking data. Movements are probably governed by several factors, some still not studied, but data show that food, water temperature, bottom type, and magnetic gradient play major roles in a shark's decision of where and when to swim. A few species exhibit differences in behavior between groups of sharks within the same geographical area. This interesting finding warrants further research to evaluate the causes of these apparent differences and whether these groups constitute different subpopulations of the same species. The lack of telemetry data on batoids and some orders of sharks must be addressed before we can gain a more comprehensive understanding of the behavior of elasmobranch fishes. Previously unpublished data from 47 smaller and 38 larger juvenile lemon sharks, collected over the decade 1988–1998, provide new results on movement patterns, habitat selection, activity rhythms, swimming speed, rate-of-movement, and homing behavior. From these results we conclude that the lemon shark is an active predator with a strong, apparently innate homing mechanism. This species shows ontogenetic differences in habitat selection and behavior, as well as differences in movements between groups of individuals within the same area. We suggest three hypotheses for future research on related topics that will help to understand the enigmatic behavior of sharks.     

Stereotypic behavior in wild marine carnivores?

Stereotypic behavior is observed in many species within zoological institutions. Attempts to reduce such behavior typically involve some form of environmental enrichment that provides opportunities for species appropriate behavior or some degree of control within the environment. However, environmental enrichment has never been completely successful in eliminating stereotypic behavior for an entire group of animals within a zoological facility. In the wild, stereotypic behavior is rarely observed. Documenting the occurrence of stereotypic behavior in the wild, and circumstances in which it occurs, could help provide insight into the causes of such behavior within zoological institutions. The following commentary details the observations of wild lemon sharks (Negaprion brevirostris) engaging in a stereotyped swimming pattern behind a research vessel north of Grand Bahama Island, Bahamas. We consider a possible explanation for the sharks' behavior and hope to stimulate conversation as well as increase examination of animal management routines in zoological facilities. Zoo Biol 30:365–370, 2011. © 2010 Wiley‐Liss, Inc.     

Entrainment of the activity rhythm of the mole crab Emerita talpoida

The mole crab Emerita talpoida migrates with the tide in the swash zone of sand beaches. A circatidal rhythm in vertical swimming underlies movement, in which mature male crabs show peak swimming activity 1-2 h after the time of high tides at the collection site. In addition, there is a secondary rhythm in activity amplitude, in which crabs are maximally active following low amplitude high tides and minimally active following high amplitude high tides. The present study determined the phase response relationship for entrainment of the circatidal rhythm with mechanical agitation and whether the cycle in activity related to tidal amplitude could be entrained by a cycle in the duration of mechanical agitation at the times of consecutive high tides. After entrainment with mechanical agitation on an orbital shaker, activity of individual crabs was monitored in constant conditions with a video system and quantified as the number of ascents from the sand each 0.5 h. Mechanical agitation at the times of high tide, mid-ebb and low tide reset the timing of the circatidal rhythm according to the timing relationship to high tide. However, mechanical agitation during flood tide had no entrainment effect. In addition, a cycle in duration of mechanical agitation entrained the rhythm in activity amplitude associated with tidal amplitude. Both rhythms and entrainment effectiveness over the tidal cycle may function to reduce the likelihood of stranding above the swash zone.     

Activity‐specific metabolic rates for diving,transiting, and resting at sea can be estimated from time–activity budgets in free‐ranging marine mammals

Time and energy are the two most important currencies in animal bioenergetics. How much time animals spend engaged in different activities with specific energetic costs ultimately defines their likelihood of surviving and successfully reproducing. However, it is extremely difficult to determine the energetic costs of independent activities for free‐ranging animals. In this study, we developed a new method to calculate activity‐specific metabolic rates, and applied it to female fur seals. We attached biologgers (that recorded GPS locations, depth profiles, and triaxial acceleration) to 12 northern (Callorhinus ursinus) and 13 Antarctic fur seals (Arctocephalus gazella), and used a hierarchical decision tree algorithm to determine time allocation between diving, transiting, resting, and performing slow movements at the surface (grooming, etc.). We concomitantly measured the total energy expenditure using the doubly‐labelled water method. We used a general least‐square model to establish the relationship between time–activity budgets and the total energy spent by each individual during their foraging trip to predict activity‐specific metabolic rates. Results show that both species allocated similar time to diving (~29%), transiting to and from their foraging grounds (~26–30%), and resting (~8–11%). However, Antarctic fur seals spent significantly more time grooming and moving slowly at the surface than northern fur seals (36% vs. 29%). Diving was the most expensive activity (~30 MJ/day if done non‐stop for 24 hr), followed by transiting at the surface (~21 MJ/day). Interestingly, metabolic rates were similar between species while on land or while slowly moving at the surface (~13 MJ/day). Overall, the average field metabolic rate was ~20 MJ/day (for all activities combined). The method we developed to calculate activity‐specific metabolic rates can be applied to terrestrial and marine species to determine the energetic costs of daily activities, as well as to predict the energetic consequences for animals forced to change their time allocations in response to environmental shifts.     

Respiration and motor activity of cercariae of three trematode species from the intertidal mollusc Littorina littorea L (Gastropoda) from the White sea

The age dynamics in oxygen consumption of the cercariae Himasthla elongata, Cryptocotyle lingua and Cercaria parvicaudata (Renicola sp.) was studied using modified Winkler's method. It was detected that under stable temperature and water salinity conditions the rate of oxygen uptake depends directly on cercariae size. The highest intensity of energetic metabolism was recorded in the first few hours of cercariae life when their movement activity was maximal. The following reduction of oxygen consumption passed unequally in three cercariae species studied. Large, long-lived H. elongata cercariae after relatively short period of active swimming turn to crawling on the bottom. The rate of oxygen uptake in such cercariae was two times less than in free-swimming ones and remained approximately invariable up to the cercariae death. The smaller sized, short-lived Renicola sp. cercariae swim actively in the water and during this time the level of their energetic metabolism remains more or less stable. It decreased drastically after cercariae sinking to the bottom, after that they perished very soon. Also relatively small C. lingua cercariae alternate the active and passive phases of swimming. Thanks to that they consume the energetic resources economically and prolong their longevity. During free-swimming period the rate of oxygen uptake of C. lingua cercariae remains more or less stable. As in the case of Renicola sp. cercariae, it decreased drastically after the cercariae sinking to the bottom. Apparently such cercariae lose their ability to infect the second intermediate host (fish).     

Behavior of the mayotte lemur,Lemur fulvus mayottensis,in captivity

A group of one male and two female Mayotte lemurs was observed at Chester Zoo, and their behaviors compared with published observations of wild populations of this and related forms of the brown lemur. It was found that the captive animals were active by both day and night, but that the pattern of activity was different from wild lemurs. The captive lemurs were less active than their wild counterparts and showed different timing of activity because of the feeding regimen at the zoo. The animals did not ignore the public at the zoo, but, on the contrary, directed some behaviors at the zoo visitors, particularly if the zoo visitors attempted to interact with the lemurs; under these conditions there was also an increase in the movements of the lemurs in the cage. Levels of aggression in the zoo lemurs were higher than those reported for wild populations, and aggression appeared to occur predominantly in the birth season. Levels of affiliative behaviors, particularly allogrooming, were comparable with those in the wild. Olfactory behaviors such as scent marking and anogenital sniffing also showed seasonal peaks and again appeared to occur at higher rates than in the wild.     

Evolution and consequences of endothermy in fishes

Regional endothermy, the conservation of metabolic heat by vascular countercurrent heat exchangers to elevate the temperature of the slow-twitch locomotor muscle, eyes and brain, or viscera, has evolved independently among several fish lineages, including lamnid sharks, billfishes, and tunas. All are large, active, pelagic species with high energy demands that undertake long-distance migrations and move vertically within the water column, thereby encountering a range of water temperatures. After summarizing the occurrence of endothermy among fishes, the evidence for two hypothesized advantages of endothermy in fishes, thermal niche expansion and enhancement of aerobic swimming performance, is analyzed using phylogenetic comparisons between endothermic fishes and their ectothermic relatives. Thermal niche expansion is supported by mapping endothermic characters onto phylogenies and by combining information about the thermal niche of extant species, the fossil record, and paleoceanographic conditions during the time that endothermic fishes radiated. However, it is difficult to show that endothermy was required for niche expansion, and adaptations other than endothermy are necessary for repeated diving below the thermocline. Although the convergent evolution of the ability to elevate slow-twitch, oxidative locomotor muscle temperatures suggests a selective advantage for that trait, comparisons of tunas and their ectothermic sister species (mackerels and bonitos) provide no direct support of the hypothesis that endothermy results in increased aerobic swimming speeds, slow-oxidative muscle power, or energetic efficiency. Endothermy is associated with higher standard metabolic rates, which may result from high aerobic capacities required by these high-performance fishes to conduct many aerobic activities simultaneously. A high standard metabolic rate indicates that the benefits of endothermy may be offset by significant energetic costs.     

Modelling energetic costs of fish swimming

The oxygen consumption rates of two cyprinid fishes, carp (Cyprinus carpio L.) and roach (Rutilus rutilus (L.)), were analysed for a wide range of body mass and swimming speed by computerized intermittent-flow respirometry. Bioenergetic models were derived, based on fish mass (M) and swimming speed (U), to predict the minimal speed and mass-specific active metabolic rate (AMR) in these fishes (AMR=aMbUc). Mass and speed together explained more than 90% of the variance in total swimming costs in both cases. The derived models show that carp consume far more oxygen at a specific speed and body mass, thus being less efficient in energy use during swimming than roach. It was further found that in carp (AMR=0.02M0.8U0.95) the metabolic increment during swimming is more strongly effected by speed, whereas in roach (AMR=0.02M0.93U0.6) it is more strongly effected by body mass. The different swimming traits of carp and roach are suitable for their respective lifestyles and ecological demands.     

COST OF TRANSPORT IN STELLER SEA LIONS, EUMETOPIAS JUBATUS

The cost of swimming is a key component in the energy budgets of marine mammals. Unfortunately, data to derive predictive allometric equations are limited, and estimates exist for only one other species of otariid. Our study measured the oxygen consumption of three juvenile Steller sea lions ( Eumetopias jubatus ) swimming in a flume tank at velocities up to 2.2 m sec⁻¹. Minimum measured cost of transport ranged from 3.5–5.3 J kg⁻¹ m⁻¹, and was reached at swimming speeds of 1.7–2.1 m s⁻¹. These cost-of-transport values are higher than those reported for other marine mammals. However, once differences in stationary metabolic rate were accounted for, the locomotor costs (LC) for the Steller sea lions were commensurate with those of other marine mammals. Locomotor costs (LC in J m⁻¹) appeared to be directly proportional to body mass (M in kg) such that LC = 1.651M^1.01. These estimates for the cost of locomotion can be incorporated into bioenergetic models and used to determine the energetic consequences of observed swimming behavior in wild marine mammals.