The cost of locomotion in North Atlantic right whales (<italic>Eubalaena glacialis</italic>)

Abstract

<p>Locomotion in any environment requires the use of energy to overcome the physical</p><p>forces inherent in the environment. Most large marine vertebrates have evolved</p><p>streamlined fusiform body shapes to minimize the resistive force of drag when in</p><p>a neutral position, but nearly all behaviors result in some increase in that force.</p><p>Too much energy devoted to locomotion may reduce the available surplus necessary</p><p>for population-level factors such as reproduction. The population of North Atlantic</p><p>right whales has not recovered following legal protection due to decreased fecundity,</p><p>including an increase in the intercalf interval, an increase in the years to first calf and</p><p>an increase in the number of nulliparous females in the population. This reproductive</p><p>impairment appears to be related to deficiencies in storing enough energy to meet the</p><p>costs of reproduction. The goal of this study was to determine whether increases in</p><p>moving between prey patches at the cost of decreased foraging opportunities could</p><p>shift these whales into a situation of negative energy gain. The first step is to</p><p>understand the locomotor costs for this species for the key behaviors of traveling and</p><p>foraging.</p><p>This study investigated the cost of locomotion in right whales by recording the</p><p>submerged diving behaviors of free-ranging individuals in both their foraging habitat</p><p>in the Bay of Fundy and their calving grounds in the South Atlantic Bight with a</p><p>suction-cupped archival tag. The data from the tags were used to quantify the oc-</p><p>currence of different behaviors and their associated swimming behaviors and explore</p><p>three behavioral strategies that reduce locomotor costs. First, the influence that</p><p>changes in blubber thickness has on the buoyancy of these whales was investigated</p><p>by comparing the descent and ascent glide durations of individual whales with differ-</p><p>ent blubber thicknesses. Next, the depth of surface dives made by animals of different</p><p>sizes was related to the depth where additional wave drag is generated. Finally, the</p><p>use of intermittent locomotion during foraging was investigated to understand how</p><p>much energy is saved by using this gait. The final piece in this study was to deter-</p><p>mine the drag related to traveling and foraging behaviors from glides recorded by</p><p>the tags and from two different numerical simulations of flow around whales. One, a</p><p>custom developed algorithm for multiphase flow, was used to determine the relative</p><p>drag, while a second commercial package was used to determine the absolute mag-</p><p>nitude of the drag force on the simplest model, the traveling animal. The resulting</p><p>drag estimates were then used in a series of theoretical models that estimated the</p><p>energetic profit remaining after shifts in the occurrence of traveling and searching</p><p>behaviors.</p><p>The diving behavior of right whales can be classified into three stereotyped be-</p><p>haviors that are characterized by differences in the time spent in different parts of the</p><p>water column. The time budgets and swimming movements during these behaviors</p><p>matched those in other species, enabling the dive shapes to be classified as foraging,</p><p>searching and traveling behaviors. Right whales with thicker blubber layers were</p><p>found to perform longer ascent glides and shorter descent glides than those with</p><p>thinner blubber layers, consistent with the hypothesis that positive buoyancy does</p><p>influence their vertical diving behavior. During horizontal traveling, whales made</p><p>shallow dives to depths that were slightly deeper than those that would cause ad-</p><p>ditional costs due to wave drag. These dives appear to allow whales to both avoid</p><p>the costs of diving as well as the costs of swimming near the surface. Next, whales</p><p>were found to glide for 12% of the bottom phases of their foraging dives, and the</p><p>use of `stroke-glide' swimming did not prolong foraging duration from that used by</p><p>continuous swimmers. Drag coefficients estimated from these glides had an average</p><p>of 0.014 during foraging dives and 0.0052 during traveling, values which fall in the</p><p>range of those reported for other marine mammals. One numerical simulation deter-</p><p>mined drag forces to be comparable, while the other drastically underestimated the</p><p>drag of all behaviors. Finally, alterations to the behavioral budgets of these animals</p><p>demonstrated their cost of locomotion constitutes a small portion (8-12%) of the</p><p>total energy consumed and only extreme increases in traveling time could result in a</p><p>negative energy balance. In summary, these results show that locomotor costs are no</p><p>more expensive in this species than those of other cetaceans and that when removed</p><p>from all the other stressors on this population, these whales are not on an energetic</p><p>`knife edge'.</p>