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Feeding Strategies and Tactics

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Feeding Strategies and Tactics 414 Feeding Strategies and Tactics Woodward, B. L., and Winn, J. P. (2006a). Apparent lateralized behav­tend to aggregate in areas with high food concentrations. For example, ior in gray whales feeding off the central British Columbia coast. Qre highest densities of polar bear s( Ursus marilimus) are found along Mar. Mamm. Sci. 22, 64-73. floe-edges and on moving ice, habitats that contain the highest den­ sity of seals: resident MUer whales{Orcintis orca) are most abundant in Johnston Strait, British Columbia when salmon migrate through the strait Also, Qre disLribuQon of humpback whales in Qre Grrlf of Mairre appears to reflect the availability of fish prey, and humpback whale distribution in southeast Alaska may partially be determined by krill Feeding Strategies and Tacticsabundance. When there are a variety of habitats available to marine rrrarrrrrrals irr a restricted area (such as rrearshore errvirorrrrrerrts), a M i c h a e l R. Heithaus and Lawrence M. D ill theoretical model predicts that, if the main concern of the animals is to maximize energy irrtake, they should be distributed proportional to the amount of food available in each habitat (Tregenza, 1995). Testing I. Introduction Qris hypoQresis is diffrcirll sirrce rrrarirre rrrarrrrnal prey availability is arine mammals are found in a wide range of habitats often hard to quanhfy. However, the distribution of botQenose dol­ including the open ocean, coastal waters, rivers, lakes, andphins (Tursiops spp.) in Shark Bay, Western Australia, conforms to M even on ice floes and land. They feed on a variety of preythis hypothesis and matches that of their fish prey in winter months species from aquatic plants to microscopic zooplankton to the largestat scales of 100s of rrrelers to kilorrreters. Similarly, Hawaiian spinner marine mammals, and a diverse array of strategies and tactics is used dolphins(Stenella longirostris) appear to match the availability of their to locate and capture these prey. Some marine mammals consumevertically migrahng prey at scales of 20 m to kilometers. Humpback and huge numbers of prey items at a time (batch feeding) while oth­ minke (Balaenoptera aciitostrata) whales do not appear’ to conform to ers attack and consume prey items singly (raptorial feeding). ManyQris hypoQresis. Instead, Qrey appear’ to show a Qrreshold resporrse to marine mammals forage in large groups while others feed alone. Inprey availability, only using a habitat once prey density has reached a this chapter, we will consider the wide range of marine mammal for­ parQcirlar level, but above this threshold there is a Qght relationship aging behaviors and the circumstances and habitats that led to the between zooplankton abundance and whale abundance. Of course, adoption of particular feeding strategies and tactics. prey availability is rrot the only factor that might irrfluence habitat use Before embarking upon a review of marine mammal foraging, it isof marine mammals, and this will be considered in detail later. important to make a distinction between a strategy and a tactic, terms Many marine mammals forage over great distances, and they may which have specific meanings in the field of Behavioral Ecolog)'. Tohave hmited knowledge of the distribution of prey patches, especially in put simply, a strategy is a genetically based decision rule (or set pelagicof habitats. In Qrese situaQorrs, marine rrrarrrrrrals, irrcludirrg pirrrri- rules) that results in the use of particulai’ tactics. Tactics are used topeds and cetaceans, may adopt movement tactics that should maximize pursue a strategy and include behaviors (Gross, 1996). Tachcs may thebe probability of encountering prey. Displacement rates are relahvely fixed or flexible, in the latter case they depend on the condition of thehigh, and movements relatively Hnear, in areas of low prey abundance, individual or characteristics of the prey or environment. For example,but arrinrals exlribit low displacerrrerrl rates arrd Iriglr turrrirrg rates (“area a hum pback whale s {Megaptera novaeangUae) strategy may be to use restricted searches”) when they encounter rich patches. that tactic which will maximize energy intake at any particular time. The whale may pursue this strategy by switching between the tactics used to capture fish and those used to catch kriU, depending upon the B. Migration relative abundance of these two prey types. When suitable habitats for a marine mammal are widely spaced, Our understanding of marine mammal foraging is hampered bymovements between them are considered migrations. There is thus the difficulty of studying these animals. They live in an environmenta continuum between habitat use decisions and migrations. Some where observations are difficult (often beneath the surface), our pres­migrations appear to be driven primarily by variation in food availa­ ence can disturb their foraging behavior, and feeding events oftenbility. Unhke baleen whales, sperm whales {Physeter macrocephalus) occur quickly and are easy to miss. Despite this, and thanks to manycannot fast for long periods of Qme, and female groups use migra- emerging technologies, much is known. We wiU begin our review byQons up to 1100 km as part of a strategy for surviring in a variable considering ways that marine mammals find and capture their prey,habitat with low local food abundance and poor foraging success. continue with a discussion of group foraging, then conclude withIn fact,a this tacQc may be the reason that female sperm whales are discussion of the causes of variation in feeding strategies and tactics.found in permanent social groups as they may benefit from the expe­ rience of old females during migrations. II. Finding Prey Migration frequendy involves trade-offs between feeding and another factor, like reproduction. Baleen whales and some pinnipeds The first step in foraging is locating prey. This may be done over many temporal and spatial scales and can involve migrations of thou­feed only for a relatively short period of time in high productivity high latitude waters, then fast during the rest of the year while moving to, sands of kilometers or switching between habitats separated by only and spending time at, low latitude breeding grounds. For example, a few meters to forage in prey-rich locations. Then, once a marine m ammal is in a prey-rich area it still m ust locate prey. northern elephant seals{Mirounga angustirostris) forage along the entire North Pacific, then migrate to a few beaches on the California coast to breed and molt. Also, some humpback whales in the Pacific A. Habitat Use Ocean reproduce in warm, low productivity, Hawaiian waters, then One way that marine mammals can increase their chances of move to the more productive waters of the north Pacific to feed dur­ encountering prey is to spend time foraging in those habitats with highing the summer months. However, some humpback whales remain in prey abundance. There is evidence that a variety of marine mammalsthe southeast Alaska feeding grounds yearround, and individuals that DWH-AR0005543 Feeding Strategies and Tactics 415 do not consume enough prey during the feeding season may forego D. Prey Detection migration to continue feeding. Marine mammals have many ways to detect their prey including vision, various types of mechanoreception, echolocation, and hear­ C. Searching and Diving ing. Most marine mammals appear to rely on vision to at least some extent. The large, forward pointing eyes of pinnipeds suggest that The way in which an animal moves through its environment can vision is an important method for detecting prey. Even species that influence its encounter rate with prey and many animals exhibit ster­ dive to extreme depths, like the elephant seal, are capable of using eotyped search patterns. Marine mammals that forage on concen­vision to find prey in dark waters at their foraging depth. Vision trated prey may continually patrol through areas where they expect to may be less important in other taxa. For example, river dolphins encounter concentrations. For example, leopard(Hydi'urga seals lep- (Platanista gangetica) of the Indian subcontinent have eyes that are tonyx) win patrol along ice edges where deparhng and returning pen­ greatly reduced and may be mostly blind. Sea otters can use their guins congregate and killer whales patrol nearshore areas in search of forepaws to find food and discriminate prey items without the aid of seals. When groups of marine mammals forage, they often spread out vision, and many pinnipeds are found in turbid waters, making vision into widely spaced subgroups and/or move forward in a Hue abreast for- a poor method of prey detection. However, they are able to use their mahon (e.g., dusky dolphins[Lagenorhynchus obscurus], pilot whales vibrissae (whiskers) to detect prey through active touch or through [Globicephda spp.], Risso’s dolphins[Grampus griseus], bottlenose minute water movements caused by their prey. dolphins, killer whales). Spreading out in such fronts may either reduce Odontocete cetaceans have a method of prey detection not foraging competition among individuals or increase the probability that available to other marine mammals—echolocation. In controlled prey is detected so the subgroups can converge to feed. situations, odontocetes can detect relatively small objects at a consid­ Once a marine mammal has selected a habitat for foraging, it erable distance. For example, a bottlenose dolphin can detect a 7.62- must execute a strategy that will optimize its net energy intake rate, cm diameter sphere from over 100 m. However, it is still unclear often with respect to trade-offs and constraints. For a diving animal how efficient echolocation is under natural conditions. It is likely to this means that it must balance the energetic costs of diving with the be less efficient than suggested by laboratory and controlled experi­ energetic gains of foraging. The costs of diving vary greatly among ments (as has been shown for bats), and may vary greatly depending marine mammals. Polar hears, sea otters[Enhydra lutris), and most on environmental conditions such as noise.
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