Whales, Whaling, and Ocean Ecosystems

Whales, Whaling, and Ocean Ecosystems

JAMES A. ESTES
DOUGLAS P. DEMASTER
DANIEL F. DOAK
TERRIE M. WILLIAMS
ROBERT L. BROWNELL
Copyright Date: 2006
Edition: 1
Pages: 418
https://www.jstor.org/stable/10.1525/j.ctt1ppsvh
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    Whales, Whaling, and Ocean Ecosystems
    Book Description:

    This unprecedented volume presents a sweeping picture of what we know about the natural history, biology, and ecology of whales in the broad context of the dynamics of ocean ecosystems. Innovative and comprehensive, the volume encompasses multiple points of view to consider the total ecological impact of industrial whaling on the world's oceans. Combining empirical research, ecological theory and modeling, and historical data, its chapters present perspectives from ecology, population biology, physiology, genetics, evolutionary history, ocean biogeography, economics, culture, and law, among other disiplines. Throughout, contributors investigate how whaling fundamentally disrupted ocean ecosystems, examine the various roles whales play in food webs, and discuss the continuing ecological chain reactions to the depletion of these large animals. In addition to reviewing what is known of the current and historic whale populations,Whales, Whaling, and Ocean Ecosystemsconsiders how this knowledge will bear on scientific approaches to conservation and whaling in the future and provocatively asks whether it is possible to restore ocean ecosystems to their pre-whaling condition.

    eISBN: 978-0-520-93320-0
    Subjects: Aquatic Sciences

Table of Contents

  1. Front Matter
    (pp. i-iv)
  2. Table of Contents
    (pp. v-vi)
  3. LIST OF CONTRIBUTORS
    (pp. vii-viii)
  4. LIST OF TABLES
    (pp. ix-x)
  5. LIST OF FIGURES
    (pp. xi-xvi)
  6. ONE Introduction
    (pp. 1-4)
    JAMES A. ESTES

    Overharvesting has led to severe reductions in the abundance and range of nearly every large vertebrate species that humans have ever found worth pursuing. These megafaunal reductions, dating in some cases from first contact with early peoples (Martin 1973), are widely known. In contrast, remarkably little is known about the ecological consequences of megafaunal extirpations. Whales and whaling are part of that legacy. Most people know that large whales have been depleted, but little thought has been given to how the depletions may have influenced ocean ecosystems. This volume is an exploration of those influences.

    My own interest in the...

  7. BACKGROUND

    • TWO Whales, Interaction Webs, and Zero-Sum Ecology
      (pp. 7-13)
      ROBERT T. PAINE

      Food webs are inescapable consequences of any multispecies study in which interactions are assumed to exist. The nexus can be pictured as links between species (e.g., Elton 1927) or as entries in a predator by prey matrix (Cohen et al. 1993). Both procedures promote the view that all ecosystems are characterized by clusters of interacting species. Both have encouraged compilations of increasingly complete trophic descriptions and the development of quantitative theory. Neither, however, confronts the issue of what constitutes a legitimate link (Paine 1988); neither can incorporate the consequences of dynamical alteration of predator (or prey) abundances or deal effectively...

    • THREE Lessons from Land Present and Past Signs of Ecological Decay and the Overture to Earth’s Sixth Mass Extinction
      (pp. 14-26)
      C. JOSH DONLAN, PAUL S. MARTIN and GARY W. ROEMER

      We are currently experiencing the sixth major extinction event in the world′s history (Thomas et al. 2004). This event is more pervasive than the previous five and is overwhelmingly human-driven. Nevertheless, distinguishing between the proximate and ultimate causes of extinction is often difficult (Caughley 1994). Alongside these extinctions, we have recently witnessed a number of complex species interactions that have restructured entire ecosystems and contributed to the decline of biodiversity. These interactions often involve apex predators, suggesting that species of high trophic status play important roles in ecosystem function (Estes 1995; Terborgh et al. 1999). This putative role of predators...

    • FOUR When Ecological Pyramids Were Upside Down
      (pp. 27-37)
      JEREMY B. C. JACKSON

      The systematic destruction of the great whales was a stupendous act of modern ecological folly that rivals the extirpation of the once vast herds of American bison in efficiency, speed, and last-minute second thoughts that marginally spared the species (Isenberg 2000; Clapham and Baker 2002; Kareiva et al., Chapter 30 of this volume; Pfister and DeMaster, Chapter 10 of this volume). It is therefore remarkable that so little serious attention has been paid until quite recently to the ecological consequences of the relatively sudden removal of whales in terms of their past roles both as keystone consumers in food chains...

    • FIVE Pelagic Ecosystem Response to a Century of Commercial Fishing and Whaling
      (pp. 38-49)
      TIMOTHY E. ESSINGTON

      The dominant role of humans as structuring agents in ecosystems is now widely acknowledged (Vitousek et al. 1997). Human activity has led to widespread alteration of marine coastal ecosystems through changes in biogeochemical cycling of nutrients, carbon and contaminants, and hydrologic regimes and through the direct and indirect effects accompanying the targeted removal of marine life. Jackson et al. (2001) conclude that human exploitation and concomitant changes in food web structure is primary among the anthropogenic alterations of coastal ecosystems. These anthropogenic alterations of community and ecosystem structure are now well documented for a diversity of ecosystems, including coral reef...

    • SIX Evidence for Bottom-Up Control of Upper-Trophic-Level Marine Populations Is It Scale-Dependent?
      (pp. 50-64)
      GEORGE L. HUNT JR.

      Understanding mechanisms of population control has been an area of concern since at least the time of Malthus, and early work in ecology emphasized the importance of energy flow for the trophic structure of ecosystems (Lindeman 1942; Odum and Odum 1955; Hutchinson 1959; Slobodkin 1960, 1962). Beginning in the 1960s, the focus was on assessing the importance of resource limitation (bottom-up mechanisms) and predation (top-down mechanisms) for population control and the structure of terrestrial (e.g., Hairston et al. 1960), lake (e.g., Brooks and Dodson 1965; Smith 1969; Zaret and Paine 1973), and rocky intertidal and subtidal ecosystems (e.g., Paine 1966,...

  8. WHALES AND WHALING

    • SEVEN Evolutionary Patterns in Cetacea Fishing Up Prey Size through Deep Time
      (pp. 67-81)
      DAVID R. LINDBERG and NICHOLAS D. PYENSON

      Food webs are dominant features of ecosystems, and predatorprey interactions are important linkages between the species that populate them. Community-level structuring forces are often invoked to explain food web dynamics, and these forces have been characterized as predominantly being either bottom-up or top-down processes (Hunter and Price 1992; Power 1992). In marine systems, the bottom-up perspective focuses on how resources and physical factors (e.g., light, nutrients, and temperature) influence food webs, beginning with phytoplankton and proceeding up through higher trophic levels. In contrast, a top-down view focuses on the influence of predators and follows their impact down to lower trophic...

    • EIGHT A Taxonomy of World Whaling Operations and Eras
      (pp. 82-101)
      RANDALL R. REEVES and TIM D. SMITH

      Whaling ranks along with some pelagic marine fishing as the world′s most spatially extensive form of exploitation of wild living resources. An understanding of its history is therefore important for analyzing the role of humans in modifying marine ecosystems.

      Whaling has involved most of the 14 mysticete (baleen) species, many of the 28 or so medium- to large-sized odontocetes (toothed whales), and numerous geographically distinct populations of these species (at least dozens). The scale of world whaling has been global, spanning bays and gulfs, continental and island shelves, and pelagic waters. Whaling began in antiquity (more than a thousand years...

    • NINE The History of Whales Read from DNA
      (pp. 102-115)
      STEPHEN R. PALUMBI and JOE ROMAN

      Of what value is knowledge about the history of a threatened species? Is it possible to chart the future of a species without information about its past? Or is knowledge about past population sizes and carrying capacities crucial to future management plans? These issues are particularly relevant to the management of populations of the great whales.

      Populations of all the baleen whales were dramatically reduced by whaling, and unprecedented international cooperation has established a global moratorium on the hunting of whales for commercial purposes until stocks recover. But what does recovery entail? For gray whales that migrate along the western...

    • TEN Changes in Marine Mammal Biomass in the Bering Sea/Aleutian Islands Region before and after the Period of Commercial Whaling
      (pp. 116-133)
      BETE PFISTER and DOUGLAS P. DEMASTER

      The Bering Sea is one of the most productive and diverse ecosystems in the world, supporting approximately 266 species of phytoplankton, 300 species of zooplankton, 450 species of fish and invertebrates, and 38 species of seabirds (Loughlin et al. 1999). Currently, 26 species of marine mammals, including large whales, beaked whales, small cetaceans, pinnipeds, the polar bear (Ursus maritimus), and the sea otter (Enhydra lutris), inhabit the area (National Research Council 1996). Many of these species were severely impacted by large-scale commercial harvesting from the mid-1700s to the mid-1900s (Clapham and Baker 2002; Webb 1988). During this time period, the...

    • ELEVEN Industrial Whaling in the North Pacific Ocean 1952–1978 Spatial Patterns of Harvest and Decline
      (pp. 134-144)
      ERIC M. DANNER, MATTHEW J. KAUFFMAN and ROBERT L. BROWNELL JR.

      For many species of conservation concern, efficient management, conservation, and harvesting strategies require information about population spatial structure. Ecologists are increasingly aware that species rarely exist as large, well-mixed populations, but rather as multiple subpopulations separated by geography and connected by various levels of dispersal. Compared to more easily studied marine mammals, relatively little is known about the population spatial structure of commercially exploited whale species (Hoelzel 1991). Accurate characterization of large regional populations as sets of smaller subpopulations can improve the accuracy of harvesting quotas (Jonzen et al. 2001), population monitoring efforts (Shelden et al. 2001), spatial location of...

    • TWELVE Worldwide Distribution and Abundance of Killer Whales
      (pp. 145-162)
      KARIN A. FORNEY and PAUL R. WADE

      Killer whales have long been recognized as formidable predators in virtually all of the world′s oceans (Leatherwood and Dahlheim 1978; Heyning and Dahlheim 1988). They are known to be common in many coastal areas, particularly at high latitudes, but they also occur in offshore and tropical waters (Leatherwood and Dahlheim 1978). Seasonal movement patterns, often associated with increased availability of prey species (Dahlheim 1981), have been documented or suggested in the eastern North Pacific (Braham and Dahlheim 1982; Baird and Dill 1995), the coast of Japan (Kasuya 1971), the North Atlantic and Norwegian waters (Sigurjónsson and Leatherwood 1988; Similä et...

    • THIRTEEN The Natural History and Ecology of Killer Whales
      (pp. 163-173)
      LANCE G. BARRETT-LENNARD and KATHY A. HEISE

      In his landmark book on marine mammals of the northeastern Pacific, whaling captain Charles Scammon remarked that killer whales ″may be regarded as marine beasts, that roam over every ocean; entering bays and lagoons where they spread terror and death among mammoth balaenas and the smaller species of dolphins, as well as pursuing the seal and walrus, devouring, in their marauding expeditions up swift rivers, numberless salmon or other large fishes that may come in their way″ (Scammon 1874:). Scammon′s observations reflect the tendency, until very recent times, of most observers to describe killer whales—and other predators—from the...

    • FOURTEEN Killer Whales as Predators of Large Baleen Whales and Sperm Whales
      (pp. 174-188)
      RANDALL R. REEVES, JOEL BERGER and PHILLIP J. CLAPHAM

      The position of the killer whale(Orcinus orca)at the top of the marine trophic pyramid is unquestioned. It consumes a remarkable variety of organisms, ranging in size from small schooling fish to blue whalesBalaenoptera musculus)and as taxonomically diverse as seabirds (ducks and alcids; Bloch and Lockyer 1988), marine reptiles (leatherback turtles,Dermochelys coriacea;Caldwell and Caldwell 1969), elasmobranchs (Fertl et al. 1996; Visser 1999a), and terrestrial mammals as they swim across coastal channels (e.g., cervids; Dahlheim and Heyning 1999). If killer whales have any natural predators, these would be other killer whales, as there is some evidence...

  9. PROCESS AND THEORY

    • FIFTEEN Physiological and Ecological Consequences of Extreme Body Size in Whales
      (pp. 191-201)
      TERRIE M. WILLIAMS

      The single most defining physical characteristic of whales in general and mysticete whales in particular is their great size. Among the odontocetes and mysticetes there is a 2000-fold range in body mass from the 50-kg vaquita(Phocoena sinus)to the 100,000-kg blue whale(Balaenoptera musculus). The smallest odontocete is 22,000 times larger than the smallest terrestrial mammal, the Etruscan shrew(Suncus etruscus), and of the 13 recognized species of mysticete (baleen) whales the smallest species, including the dwarf minke whale(Balaenoptera acutorostratasubspecies)and pygmy right whale(Caperea marginata), are nearly 10 times larger than the biggest flesh-eating terrestrial mammals...

    • SIXTEEN Ecosystem Impact of the Decline of Large Whales in the North Pacific
      (pp. 202-214)
      DONALD A. CROLL, RAPHAEL KUDELA and BERNIE R. TERSHY

      Biodiversity loss can significantly alter ecosystem processes (Chapin et al. 2000), and ecological extinction can have similar effects (Jackson et al. 2001). For marine vertebrates, overharvesting is the main driver of ecological extinction, and the expansion of fishing fleets into the open ocean has precipitated rapid declines in pelagic apex predators such as whales (Baker and Clapham 2002), sharks (Baum et al. 2003), tuna, and billfishes (Cox et al. 2002; Christensen et al. 2003), leading to a trend in global fisheries toward exploitation of lower trophic levels (Pauly et al. 1998a). Globally, many fish stocks are overexploited (Steneck 1998), and...

    • SEVENTEEN The Removal of Large Whales from the Southern Ocean Evidence for Long-Term Ecosystem Effects?
      (pp. 215-230)
      LISA T. BALLANCE, ROBERT L. PITMAN, ROGER P. HEWITT, DONALD B. SINIFF, WAYNE Z. TRIVELPIECE, PHILLIP J. CLAPHAM and ROBERT L. BROWNELL JR.

      The Southern Ocean can broadly be defined as the oceanic region surrounding the continent of Antarctica and south of the Antarctic Convergence, or Polar Front (Figure 17.1A). Because of its latitude, it is an ecosystem of extreme seasonal variability. This variability is ultimately related to sunlight—its presence in the southern summer, and absence in the winter, results in dramatic changes in the extent of sea ice around the continent (Figure 17.1B, C), in the degree of primary productivity of the ocean, and in the distribution, abundance, and life history strategies of all organisms that live there.

      Among these organisms...

    • EIGHTEEN Great Whales as Prey Using Demography and Bioenergetics to Infer Interactions in Marine Mammal Communities
      (pp. 231-244)
      DANIEL F. DOAK, TERRIE M. WILLIAMS and JAMES A. ESTES

      An increasing difficulty in many areas of conservation is the need to formulate and test theories about biological processes and management outcomes that occur over vast areas. Unfortunately, these large-scale problems are poorly matched with most research done in the name of conservation. Academic, and even nonacademic, conservation research typically focuses on one or a few species, often with detailed data, and usually at small scales, even as the scale of decision making and management undertaken by government agencies and nongovernmental organizations (NGOs) becomes ever larger. While many have decried this divorce between real-world conservation problems and conservation research, the...

    • NINETEEN Whales and Whaling in the North Pacific Ocean and Bering Sea Oceanographic Insights and Ecosystem Impacts
      (pp. 245-261)
      ALAN M. SPRINGER, GUS B. VAN VLIET, JOHN F. PIATT and ERIC M. DANNER

      Hideo Omura′s concern over the killing of fin whales in the early 1950s, just a few years after the resumption of whaling following World War II, foreshadowed the demise of most of the remaining great whales in the following two decades as the slaughter expanded across the North Pacific (Springer et al. 2003). His concern was warranted because of the development of an unprecedented human capability for large-scale harvests of even the fastest whales using high-speed catcher boats and mechanized factory ships designed specifically for this purpose. In 1955, the nominal harvest of fin whales was about 2,100, and it...

    • TWENTY Legacy of Industrial Whaling Could Killer Whales Be Responsible for Declines of Sea Lions, Elephant Seals, and Minke Whales in the Southern Hemisphere?
      (pp. 262-278)
      TREVOR A. BRANCH and TERRIE M. WILLIAMS

      Industrial whaling decimated populations of the great whales in the Southern Hemisphere and undoubtedly had a marked effect on the ecosystems in which they existed. In the Southern Hemisphere, industrial whaling fleets focused on a succession of less and less valuable species, from humpback(Megaptera novaeangliae), blue(Balaenoptera musculus), fin(B. physalus), sperm(Physeter macrocephalus), and sei whales(B. borealis)to, finally, Antarctic minke whales(B. acutorostrata)(Hilborn et al. 2003). This pattern occurred after the decimation of southern right whales(Eubalaena glacialis australis)in the nineteenth century. As a result, by the early 1970s, only Antarctic minke whales remained...

    • TWENTY-ONE Predator Diet Breadth and Prey Population Dynamics Mechanism and Modeling
      (pp. 279-285)
      MARC MANGEL and NICHOLAS WOLF

      The notion that the sequential megafaunal collapse of several subpopulations of harbor seal, Steller sea lion, and sea otter (sensu Springer et al. 2003) is due to killer whale predation relies on the assumption of expanding diet breadth for orcas. That is, the putative mechanism for the decline is a preference of killer whale for large whale species, but an inclusion of certain populations of harbor seal, fur seal, Steller sea lion, and sea otter as the preferred prey types become less available. When considering the observed sequential declines, a number of features need to be explained (Figure 2 of...

    • TWENTY-TWO Bigger Is Better The Role of Whales as Detritus in Marine Ecosystems
      (pp. 286-300)
      CRAIG R. SMITH

      Organic detritus (i.e., nonliving organic matter) plays fundamental roles in the structure and dynamics of all marine ecosystems. The importance of a particular type of organic detritus in an ecosystem depends on several key characteristics of the material, including (1) the size of the detrital particles; (2) the nature of organic materials contained within the particles (e.g., the presence of nutritious lipids and proteins); (3) the flux of organic carbon, or limiting nutrient, entering the ecosystem in the form of the detritus (especially relative to fluxes in other forms); (4) the frequency of occurrence of the detrital particles (essentially flux...

  10. CASE STUDIES

    • TWENTY-THREE Gray Whales in the Bering and Chukchi Seas
      (pp. 303-313)
      RAYMOND C. HIGHSMITH, KENNETH O. COYLE, BODIL A. BLUHM and BRENDA KONAR

      Among the large cetaceans, gray whales(Eschrichtius robustus)are unique in three important ways: They are benthic feeders; they undertake one of the longest migrations of any mammal; and they may be fully recovered (at least the eastern Pacific stock) from overharvesting by commercial whaling. The eastern (Chukotka-California) gray whales migrate annually between the mating regions and calving lagoons on the west coast of Baja California, Mexico, to summer feeding grounds in the northern Bering Sea and the Chukchi Sea (Rice and Wolman 1971; Marquette and Braham 1982; Findley and Vidal 2002). The purpose of this chapter is to explore...

    • TWENTY-FOUR Whales, Whaling, and Ecosystems in the North Atlantic Ocean
      (pp. 314-323)
      PHILLIP J. CLAPHAM and JASON S. LINK

      Although whaling for subsistence purposes has occurred in various locations worldwide from prehistoric times, commercial whaling—in the sense of a continuous, directed effort pursued for profit—likely had its origin in the North Atlantic Ocean (Figure 24.1). The earliest known commercial ventures were those conducted by the Basques of northern Spain and southern France, who operated a well-organized hunt in the Bay of Biscay beginning in or before the eleventh century (Aguilar 1986; Reeves and Smith, Chapter 8 of this volume). Various coastal whale hunts existed in Europe in medieval times, and by the sixteenth century whalers were already...

    • TWENTY-FIVE Sperm Whales in Ocean Ecosystems
      (pp. 324-334)
      HAL WHITEHEAD

      For Herman Melville the sperm whale,Physeter macrocephalus, was ″The Whale,″ and many supposedly generic illustrations of whales portray sperm whales. But the sperm whale is not a generic whale; in fact, from almost any perspective, it is the most atypical of species. One class of attributes by which the sperm whale has no close analogs, or homologs, is ecology. The ecology of the sperm whale forms the principal subject of this chapter. I will describe the trophic web surrounding the sperm whale with the goal of addressing three questions: How are sperm whale populations regulated? How do sperm whales...

    • TWENTY-SIX Ecosystem Effects of Fishing and Whaling in the North Pacific and Atlantic Oceans
      (pp. 335-343)
      BORIS WORM, HEIKE K. LOTZE and RANSOM A. MYERS

      Human alterations of marine ecosystems have occurred throughout history, but only over the last century have these reached global proportions. Three major types of changes have been described: (1) the changing of nutrient cycles and climate, which may affect ecosystem structure from the bottom up, (2) fishing, which may affect ecosystems from the top down, and (3) habitat alteration and pollution, which affect all trophic levels and therefore were recently termed side-in impacts (Lotze and Milewski 2004). Although the large-scale consequences of these changes for marine food webs and ecosystems are only beginning to be understood (Pauly et al. 1998;...

    • TWENTY-SEVEN Potential Influences of Whaling on the Status and Trends of Pinniped Populations
      (pp. 344-360)
      DANIEL P. COSTA, MICHAEL J. WEISE and JOHN P. Y. ARNOULD

      Although this volume focuses on whales and whaling, the depletion of great whales over the last 50 to 150 years perturbed the marine interaction web, thus influencing many other species and ecosystem processes (Estes, Chapter 1 of this volume; Paine, Chapter 2 of this volume). Such interaction web effects have been hypothesized for several pinniped species. For example, the reduction of great whales in the Southern Ocean may have caused seal and penguin populations to increase because of reduced competition for their shared prey, krill (Laws 1977; Ballance et al., Chapter 17 of this volume). In addition, pinnipeds share some...

  11. SOCIAL CONTEXT

    • TWENTY-EIGHT The Dynamic Between Social Systems and Ocean Ecosystems Are There Lessons from Commercial Whaling?
      (pp. 363-372)
      DANIEL W. BROMLEY

      The general issue to be explored here concerns the dynamic nexus between the gradual intensification of human interaction with—and extraction from—nature and the inevitable biological response to that activity. We may think of this dynamic as an instance of co-evolutionary adaptation in which (1) biological processes undergo transformation in the face of extensive human exploitation, and (2) human processes (and associations) in turn undergo transformation in the face of biological feedback onto human communities that have been organized and structured around this very interaction. Much of the literature in this general area concerns the first phenomena—humans bringing...

    • TWENTY-NINE Whaling, Law, and Culture
      (pp. 373-376)
      MICHAEL K. ORBACH

      All environmental law and policy, including policy governing whaling, involves trade-offs between the state of the world′sbiophysicalecology (i.e., all nonhuman elements of the world and their relationships with one another) and the state of the world′shumanecology (i.e., humans and their relationships with one another, including human governance institutions). All rules of governance directly affect only human behavior; through that behavior they shape the biophysical world. The configuration of the biophysical environment, in turn, defines the form of the costs and benefits incurred and received by humans in the use of that environment. Furthermore, every decision regarding...

  12. OVERVIEW AND SYNTHESIS

    • THIRTY Whales Are Big and It Matters
      (pp. 379-387)
      PETER KAREIVA, CHRISTOPHER YUAN-FARRELL and CASEY O’CONNOR

      Whales have a unique place in conservation lore. Their plight is widely known, and the beaching of even a single whale is a major news event, typically drawing hundreds of spectators. In addition, as a marine mammal, whales are given favored legal protection above and beyond that given to the bulk of the planet′s biodiversity. The irony is that, although whales have become a symbol of the human capacity for greedy overharvest and a rallying point for environmental activists, we know surprisingly little about their ecological role. Our ignorance regarding the ecology of whales is symptomatic of conservation: Too often...

    • THIRTY-ONE Retrospection and Review
      (pp. 388-394)
      J. A. ESTES, D. P. DEMASTER, R. L. BROWNELL JR., D. F. DOAK and T. M. WILLIAMS

      This final chapter is a review of the volume′s content, written with the intent of revisiting our initial goals (Chapter 1), assessing the degree to which we have succeeded in achieving these goals, and providing some direction for future research on the central question underlying this volume: How did whales and whaling influence the dynamics of ocean ecosystems? Our synopsis also highlights the significant findings and ideas in each of the preceding chapters.

      We begin by reiterating both the reasons for wondering about the influences of whales and whaling, and our approach to addressing the question. Given the diversity of...

  13. INDEX
    (pp. 395-402)