Whales, Whaling, and Ocean Ecosystems

F I G U R E 3 0.2. Primary and secondary pelagic consumers of krill and squid. Modeled after Laws 1985.

Bering Sea and possible prey switching by killer whales abundance recover. The expected recovery of great whales in
(Estes, Chapter 1 of this volume; Estes et al. 1998; Springer the Pacific Ocean will provide a terrific opportunity to assess
et al. 2003) is yet another line of evidence suggesting that the killer whale hypothesis.
hunting whales led to major shifts in species interactions
with reverberations throughout marine ecosystems. Certainly, as whale numbers increase, we can expect a
Although more research is needed in this area, the weight of reduction in the primary prey of many whales, such as krill;
evidence to date suggests that whales profoundly alter the the effects of which will resonate throughout the food web
composition and functioning of oceanic ecosystems. (Butman et al. 1995; Laws 1985; May et al. 1979). For exam-
ple, despite the enormous complexity of the Southern Ocean
Some Whales are Coming Back—What Will This ecosystem and the difficulty in sorting out the myriad influ-
Mean for Our Oceans? ences affecting the abundance of krill and their predators
(Ballance et al., Chapter 17 of this volume), it is hard to imag-
Any substantial increase in whale numbers is likely to have ine the recovery of baleen whales not having a dramatic
profound effects on ocean ecosystems. In fact, much of the effect on krill and the Southern Ocean food web. Krill in the
motivation for this book is the spark provided by the con- Southern Ocean are eaten primarily and secondarily by
troversy surrounding the National Marine Fisheries Service’s squid, fish, birds, seals, and whales (Figure 30.2). Since 1900,
(NMFS) conclusion that indirect fishing effects could be con- whale contribution to total annual krill consumption has
tributing to Steller sea lion declines, and therefore the pollock dropped from over 40% to less than 10%, which equates to
fishery needed to be limited (NRC 2003b). Contrary to 170 million and 35 million metric tons, respectively (Laws
NMFS’s inference, several scientists have suggested that pol- 1985). If whales increase in abundance and deplete krill pop-
lock may have no role or a negligible role in the sea lion ulations, other consumers may suffer. Moreover, as whales
declines; instead, recent declines in Steller sea lions could recover, they will compete with the commercial fleets that
have been driven by a modest number of killer whales harvest krill. The krill fishery now takes in an estimated
switching to the consumption of sea lions after having lost 160,000 metric tons annually, and demand for commercially
great whales as potential food sources through overharvest harvested krill is expected to rise in coming decades, both
(Estes et al. 2004; NRC 2003b; Springer et al. 2003; Williams within current markets and with possible uses in biotech-
et al. 2004). This scenario is at best a hypothesis, but it is a nology and pharmaceuticals (Nicol and Endo 1997).
plausible hypothesis and an idea that should be taken seri-
ously. If true, then as the larger prey of killer whales recover, In general, a return of cetacean primacy could impact the
predation on the pinnipeds should be relaxed and pinniped prey availability for many fishes currently harvested by
commercial fisheries (Essington, Chapter 5 of this volume;
Croll et al., Chapter 16 of this volume), with unanticipated

384 O V E R V I E W A N D S Y N T H E S I S

consequences. When it comes to harvest and depletion of factor such as temperature could explain a great deal of
fish or squid populations, humans have had no rivals over the annual variation in population change, yet it could be
the last century. It is interesting to speculate what it would predation that determines the persistence and average
mean for whales to resume historical abundances and them- abundance of a species (Hassell et al. 1998). Ballance et al.
selves act as major consumers. Because of their energetic (Chapter 17 of this volume) recognize this point when they
demands and adaptations for open-ocean foraging, whales remind us that all of our studies of pinnipeds, krill, and
could compete with the commercially important species for seabirds have taken place long after whales were severely
limited prey resources. According to Croll et al. (Chapter 16 depleted and that we might see an entirely different ecosys-
of this volume), current whale populations in the North tem if whales returned to historical levels.
Pacific are already comparable with commercial fisheries in
terms of consumption (55,155 metric tons day−1 and One of the best statistical explorations of top-down versus
75,468 metric tons day−1, respectively) and trophic level bottom-up hypotheses in ocean systems does not concern
(3.4 compared with 3.2, respectively). The regulation and whales but is nonetheless informative. In a study of cod-
management of fisheries has always suffered from a myopic shrimp interactions in the North Atlantic Ocean, Worm and
single-species syndrome (Zabel et al. 2003); the recovery of Myers (2003) found compelling evidence of top-down con-
whales could finally force fisheries managers to abandon the trol, mediated somewhat by temperature. The decline of cod
foolishness of managing fisheries as though harvested species (due to overharvest) has clearly led to an increase in shrimp
exist in ecological vacuums. (a common prey of cod). In contrast to much of the discus-
sion regarding correlations between climate and pinnipeds or
Which Is More Important: Whale Abundance or krill, Worm and Myers adopted a formal hypothesis-testing
Environmental Fluctuations? approach entailing competing hypotheses. Without a for-
mal contrast of competing hypotheses, spatial correlations
There has been a long-standing debate in ecology between between primary productivity and consumer abundances
those who emphasize environmental factors and those who simply do not imply an absence of top-down control, as
emphasize biotic interactions and trophic effects. These alter- some have argued (Hunt, Chapter 6 of this volume). Of
native perspectives about population regulation pervade such course, the only incontrovertible way of establishing which
subjects as density dependence (Smith 1961), supply-side is more important—top-down or bottom-up forces—is an
ecology (Lewin 1986), and the debate regarding bottom-up experiment that simultaneously manipulates both in a fac-
versus top-down control of ecosystems (Hunter and Price torial manner.
1992; see also Matson and Hunter 1992). Some researchers,
including contributors to this volume (e.g., Hunt, Chapter 6 There has been one notable meta-analysis of experimental
of this volume), discount the hypothesis that whales can manipulations performed in marine ecosystems, in which
drive marine ecosystems, because they take the observation the effect of removing predators versus adding nutrients were
that fish populations often track climate variability as evi- compared (Micheli 1999). Responses were measured in terms
dence against top-down control. It may be that the debate of the log ratio of treatment (either predator removal or nutri-
about top-down and bottom-up control is more a result of ent supplementation) to control biomass (more precisely as
the nuances of language than fundamental scientific differ- “log response ratios,” which are the log-transformed ratios of
ences. For example, Hunt (Chapter 6 of this volume) con- the treatment mean divided by the control mean, weighted
cludes that the strong responses of fish, pinnipeds, and by sampling variances). Both nutrient supplementation and
seabirds to climate variability are a “manifestation of bottom- predator removal had effects in these marine experiments,
up forcing.” That is a reasonable inference. But “forcing” is but predator removal yielded effects on average 2–3 times
different than control. The environment and food supply larger than the effects of nutrient addition. Obviously none
can drive fluctuations, while at the same time higher trophic of the experiments reviewed by Micheli involved whales, but
levels constrain the average abundances around which pop- they were marine systems with similar numbers of trophic
ulations fluctuate. levels to those found in whale food webs.

It is crucial to distinguish between those factors that best Since experiments are impractical for whales, and corre-
statistically account for variation in rates of population lations between population fluctuations and environmental
change and those factors that govern the persistence and conditions are by themselves suspect, what might be tools
long-run average abundance of a species. In the literature for testing ideas about whales as agents of top-down control
discussing whales and the food webs in which they are of marine systems? We think the most fruitful approach is
embedded, too much is made of casual correlations between an examination of mechanistic models of trophic interac-
population fluctuations and environmental variation. If one tion, with estimates of energetic needs and consumption
is to take a statistical approach when assessing top-down ver- rates (e.g., Doak et al., Chapter 18 of this volume). These
sus bottom-up factors, there must be an analysis that includes models lead to much more precise hypotheses than envi-
variation in the upper trophic levels and not simply variation ronmental correlations do, and these hypotheses can be
in environmental conditions or primary productivity. A incontrovertibly falsified by data on consumption rates,
mortality, or responses to altered predator density. Spatial
or temporal variation in whale abundance and in whale

W H A L E S A R E B I G A N D I T M A T T E R S 385

recovery can be used in conjunction with these models to Colburn, T. and M. J. Smolen. 1996. Epidemiological analysis of
test their merit. persistent orgaochlorine contaminants in cetaceans. Review of
Environmental Contamination Toxicology 146: 91–172.
Our Understanding of Ocean Ecology Should be
Enhanced Due to the Recovery of Whales Estes, J.A., E.M. Danner, D.F. Doak, B. Konar, A.M. Springer, P.D.
Steinberg, M.T. Tinker, and T.M. Williams. 2004. Complex
The recovery of whales is a huge “experiment” at the scale trophic interactions in kelp forest ecosystems. Bulletin of
of ocean subbasins. Because whale recovery will be uneven, Marine Sciences 74: 621–638.
we have an opportunity to make predictions about food-web
impacts and community impacts that could then be tested Estes, J.A., M.T. Tinker, T.M. Williams, and D.F. Doak. 1998.
over the next 50 years. But we will never learn anything Killer whale predation on sea otters linking oceanic and
from whale recovery if we neglect their ecological interac- nearshore ecosystems. Science 282: 473–476.
tions and continue to focus solely on their numbers.
Whereas intertidal and terrestrial ecologists have long Evans, P.G.H. 1987. The natural history of whales and dolphins.
recognized the importance of species interactions and the New York: Facts on File.
interplay of competition and predation, biological oceanog-
raphy has focused on primary productivity and nutrients. Flannery, T. 2001. The eternal frontier. New York: Grove Press.
When top-down effects have been looked for (Worm and Frantzis. A. 1998. Does acoustic testing strand whales? Nature
Myers 2003; Micheli 1999; Paine, Chapter 2 of this volume),
they have been found in marine systems. Studies of whale 392: 29.
ecology and recovery need to examine the possibility of pro- Fujiwara, M. and H. Caswell. 2001. Demography of the endan-
found top-down impacts. Much could be gained by simul-
taneously sampling whale populations and key components gered north Atlantic right whale. Nature 414: 537–541.
of the biotic community in ocean subbasins where whales Gerber, L.R., D.P. DeMaster, and S.P. Roberts. 2000. Measuring
are increasing and in subbasins where they are not. Of
course, this is not a real experiment because of lack of success in conservation. Scientific American 88: 316–324.
randomization and control; nonetheless, appropriate obser- Greenpeace. 2004. Whales. Accessed May 19, 2004, at http://
vational studies of contrasting ocean subbasins in conjunc-
tion with models such as those put forth by May et al. (1979) w w w. g r e e n p e a c e . o r g / i n t e r n a t i o n a l _ e n / c a m p a i g n s /
can yield strong inferences. The recovery of whales could intro?campaign_id=4017.
provide the biological perturbation that changes the way Greene, A. J. and C.H. Pershing. 2004. Climate and conservation
we think about oceans—but only if we open our eyes to biology of North Atlantic right whales: the right whale at the
species interactions and trophic effects. wrong time? Frontiers in Ecology and Environment 2: 29–34.
Hassell, M.P., M.J. Crawley, H.C.J. Godfray, and J.H. Lawton.
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T H I R T Y- O N E

Retrospection and Review

J. A. E STE S, D. P. D E MASTE R, R. L. B R OWN E LL, J R., D. F. D OAK,
AN D T. M. W I LLIAM S

This final chapter is a review of the volume’s content, written detritus. The other arose from the different approaches people
with the intent of revisiting our initial goals (Chapter 1), take in practicing science: as theory and broad concepts, as the
assessing the degree to which we have succeeded in achiev- interface across academic disciplines, and as case studies.
ing these goals, and providing some direction for future Recognizing these differences, we decided to invite two kinds of
research on the central question underlying this volume: people: those who were familiar with whales and whaling and
How did whales and whaling influence the dynamics of those who were familiar with the relevant theories or scientific
ocean ecosystems? Our synopsis also highlights the signifi- disciplines. These groups were not mutually exclusive, because
cant findings and ideas in each of the preceding chapters. most people who study whales also have broader disciplinary
interests. However, there were several participants who were
We begin by reiterating both the reasons for wondering invited because of their disciplinary skills and had little or no
about the influences of whales and whaling, and our prior knowledge of either whales or whaling.
approach to addressing the question. Given the diversity of
species, habitats, and conservation problems in today’s Why Are Whales Interesting?
world, why should whales and whaling deserve special atten-
tion? The answer at one level is simply a matter of opinion The editors of this volume were drawn to questions con-
and choice. Because of our various backgrounds, each of the cerning whales and whaling by two forces. One derives from
authors of this volume was interested in larger issues that the whales themselves. As the largest animals on earth, their
involved these topics. Different personal histories might have biology is inherently interesting. As oceanic mammals
led one or more of us in different directions, so there is an primarily hidden from man’s view, their role in ecosystems
element of serendipity in what has been produced. remains a mystery. The important organismal-level features
of whales are discussed by Williams in Chapter 15, who
Our reasons for asking about the ecological effects of whales explores the problems of being extremely large, and the phys-
and whaling were logical and multifaceted. However, the best iological and ecological consequences of large body size. This
way of addressing these questions was far less clear and required unique biology established an a priori case for the ecological
two key elements: a conceptual structure and the right people importance of whales and the subsequent impact of their
to implement it. The conceptual structure emerged in two forms. disappearance through whaling.
One of these arose from the ways we imagined that whales influ-
enced ecosystem-level processes: as predators, as prey, and as

388

Whaling, a human endeavor, substantially changed many whales are there? No one would challenge the claim
the distribution and abundance of most whale species across that the abundance of most large whale species was reduced
the world’s oceans (chronicled in many of the chapters). The by whaling. That point is made repeatedly throughout the
same is true of many other large vertebrates, but these volume and, indeed, is the fundamental event upon which
changes seemed especially important for whales because of the volume is based. However, the magnitude of these reduc-
their great historical abundance on the one hand and their tions is a matter of uncertainty and debate. Reeves and Smith
large size, high metabolic rates, and high trophic status on (Chapter 8) provide a historical synopsis of modern whaling,
the other. including its chronology, methods, species, and geography,
beginning with the Basques more than a thousand years ago
The other force that drew us together was a growing appre- and ending with the sudden curtailment of industrial-level
ciation in ecology and conservation biology for the func- commercial whaling in the 1970s. Logbook records from the
tional importance of top-down forcing processes initiated by various whale fisheries discussed by Reeves and Smith pro-
high-trophic-status consumers, a point that is emphasized in vide one basis for estimating the size of whale stocks before
various ways in Chapters 2, 3, and 4. Chapter 2, by Paine, whaling, and thus the degree to which they have been
provides an overview of both the relevant ecological concepts depleted. An alternative approach, described and evaluated
and theory, and from that view asks the question: What by Palumbi and Roman (Chapter 9), is based on theoretical
might we infer about the ecological effects of whales and relationships between genetic diversity and population size,
whaling, based on what is known about food web dynamics combined with genetic information from living whales.
from other, more tractable species and ecosystems? Paine When applied to fin and humpback whale populations in the
concludes that if photosynthetic carbon in the world’s oceans North Atlantic, this approach provides population estimates
is largely or entirely utilized, the conclusion that whales and greatly exceeding those obtained from logbook records, a
whaling were ecologically significant forces is inescapable. In conclusion that has generated a great deal of debate (IWC
Chapter 4, Donlan et al. extend Paine’s conceptual argument 2005). Palumbi and Roman argue that new analyses of log-
in two dimensions: to the land and over deep time, with an book records are unlikely to provide much additional insight
emphasis on the functional importance of megafauna. The into the different conclusions of these methods. Conversely,
authors point out that on these expansive scales of space genetic-based estimates will inevitably improve as more
and time, ecological influences by the megafauna can be detailed information (i.e., more genes sequenced from more
seen in two ways: through the loss of native species and by individuals of more species) becomes available and as the the-
the introduction of exotics. Donlan et al. reinforce the long- oretical bases for these estimates are refined. On the other
standing argument that humans have played a significant hand, those who question the veracity of the Palumbi and
role in the megafaunal demise, both recently and in the past. Roman estimates (IWC 2005) believe that a more thorough
They further suggest that these losses left many species integration of the logbook records and whale process records
anachronistic on their landscapes, thus leading in some cases and capability would likely lead to the conclusion that whale
to dysfunctional ecosystems. Chapter 5 provides a similar populations as large as those proposed by Palumbi and
view, but for the coastal oceans. In this chapter Jackson makes Roman (Chapter 9) were unlikely. Where the truth lies is not
a plea for “retrospective ecology,” arguing that quasi-pristine a mere academic curiosity, since estimates of prewhaling
modern ecosystems are time machines for understanding numbers will influence both recovery targets and the assess-
how systems worked when their megafauna were intact. He ments of ecosystem-level impacts of whaling.
explores this idea for the coastal oceans (emphasizing the
Caribbean) and concludes that the loss of large vertebrates Other chapters in this volume provide additional infor-
was of immense ecological consequence—among other mation concerning the abundance and distribution of
things, leading to destabilizations that in turn may have whales before and after whaling. By relating spatiotemporal
subsequently contributed to reduced production and carry- patterns in the North Pacific harvest records to various
ing capacity. Seen in this light, whales and their depletion oceanographic features of that region, Springer et al. (Chap-
by whaling provide a relatively recent and well-understood ter 19) identify potential oceanographic hot spots. This pro-
case of the repeated loss of megafauna around the world, vides a mechanistic basis for the significant stock structure
allowing us a better chance to think about and analyze the suggested by Danner et al. (Chapter 11) and for the related
consequences of such losses than is possible for many other implication that local rates of whale depletion and ecosys-
systems. tem effects of whaling were much greater than might be
inferred solely from an ocean basin–wide view. Likewise,
Conceptual Structure and Approaches Springer et al. point out that vastly more whales were taken
to the Question from the southwestern region, whereas bounty and preda-
tor-control programs likely had stronger depletion effects
The Question of Population Size on pinnipeds in southeast Alaska. Perhaps, then, it is not sur-
prising that sea otter and pinniped populations have col-
Central to understanding the interplay among whales, whaling lapsed in southwest Alaska, whereas this generally has not
and ocean ecosystems is the seemingly simple question: How occurred in southeast Alaska.

R E T R O S P E C T I O N A N D R E V I E W 389

Pfister and DeMaster (Chapter 10) provide another per- Highsmith et al. establish that gray whale feeding in the shal-
spective on the greater North Pacific ecosystem by contrasting low soft-sediment benthos has had important effects on this
historic and modern estimates of marine mammal biomass ecosystem by consuming epi- and infaunal organisms and
among species and species groups. Although these authors through the resuspension of sediments and nutrients from
carefully qualify their results based on many uncertainties and the benthos to the water column. These authors also point out,
the poor quality of some of the data, their findings demon- however, that such effects vary spatially, depending on such
strate that marine mammal biomass in the North Pacific was factors as sediment type and local oceanographic features.
and currently is dominated by the large whales, even though
abundances are now much reduced across most taxa. The general conclusion that large whales are (or were)
important consumers in ocean ecosystems is reinforced by
In addition to understanding the population size of great several other chapters. Essington (Chapter 5) explored the
whales, there has been much recent interest in determining larger issue of the effects of fishing and whaling on pelagic
the abundance of killer whales in ocean ecosystems. This ecosystems by using known food web structures and dynamic
stems from claims that some killer whales obtained signifi- food web models to ask how this system may have changed
cant nutritional input by feeding on large whales (Springer following population reductions of the large consumers, in
et al. 2003) or large-whale carcasses (Whitehead and Reeves this case the sperm whales, tunas, and billfishes. Essington
2005). It has further been suggested that as these nutritional concludes that population reductions of these species led to
resources declined due to whaling, killer whales may have fed changes, but these appear to be modest compared with those
more intensively on pinnipeds and sea otters, thus driving observed in other ecosystems. Whitehead (Chapter 25)
their populations downward in some areas (Chapters 1, 14, reached a similar conclusion for sperm whales, based on the
18, 20). The distribution and abundance of killer whales is an great abundance and high trophic status of this species. Con-
important component in any analysis of these hypotheses. versely, Essington reports that his modeling approach cannot
Chapter 12, by Forney and Wade, provides such a worldwide capture qualitative shifts in process; thus, the possibility
assessment of the distribution and abundance of killer of more radical shifts in ecosystem organization cannot be
whales. These authors conclude that the world killer whale discounted.
population presently exceeds 50,000 individuals, with the
greatest population density occurring at high latitudes. Inter- A different approach is taken by Croll et al. (Chapter 16).
estingly, modern-day patterns of killer whale distribution Here estimates of ocean production, whale abundance, whale
and abundance correlate well with global patterns of ocean trophic status, and trophic transfer efficiency are used to
productivity. Given the interest and controversy over the evaluate the potential impact of foraging whales in ocean
ecological importance of killer whale predation, it is appro- ecosystems. Focusing on the North Pacific (although other
priate that the Forney and Wade chapter is followed by a more regions would likely provide similar results), these authors
detailed account by Barrett-Lennard and Heise (Chapter 13) of suggest that, prior to whaling, the large whales coopted more
the natural history and behavior of killer whales. These than 50% of the total primary production, and that after
authors focus on the well known dichotomy between fish whaling this was reduced to less than 10%. Croll et al. are
eating (resident types) and marine mammal eating (transient careful to explain that these calculations do not necessarily
type) whales, pointing out that this division has long been transfer directly to the strength of food web interactions.
known, and that clear genetic differences exist between However, based on similar measurements and analyses from
resident and transient whales. Different feeding behaviors or other species and ecosystems in which consumer-mediated
dietary preferences are the products of long periods of cul- influences are known or thought to be important, Croll
tural evolution. However, a great deal of variation in diet et al.’s conclusions fall closely in line with those made by
and foraging behavior exists within each type. The authors other authors in this volume—that the great whales were
also point out an apparent paradox in killer whale diet and important consumers in ocean ecosystems, and this interac-
foraging behavior—a strong inertia against change on the tion likely was substantially altered by the effects of whaling.
one hand, and the capacity for rapid change on the other.
Focusing mainly on mysticete/krill interactions in the
Large Whales as Predators Southern Ocean, Ballance et al. (Chapter 17) also concluded
that whaling must have exerted a significant force on this
But what evidence is there for ecologically important influ- food web, although these influences cannot easily be sepa-
ences of whales, beyond their former abundance and cur- rated from the confounding effects of oceanographic change.
rently reduced numbers? To answer that question, we sought The latter in particular clearly affected krill population
contributors to address evidence for the role of large whales dynamics and, therefore, the population dynamics of krill
as important consumers. In this case there is evidence available consumers. Worm et al. (Chapter 26) approached the ques-
from each of the three scientific approaches—theoretical/ tion of whaling and fishing effects on ocean ecosystems by
conceptual, disciplinary interfaces, and specific case studies. exploring time series of whale abundance estimates and stock
The most concrete and definitive evidence comes from studies assessment data for fin fishes and forage fishes in the North
on gray whales in the Bering and Chukchi seas (Chapter 23). Atlantic and North Pacific oceans. These authors concluded
that there are significant trophic linkages among various for-
age fish, fin fishes, and great whales, and that whaling prob-

390 OVERVIEW AND SYNTHESIS

ably reduced the intensity of competition between some papers by Whitehead and Reeves (2005) and DeMaster et al.
species of whales and fin fishes for a limiting forage fish (2006).
resource. The result was increased fin fish populations, espe-
cially during the later part of the twentieth century in the There have been two proposed examples of the so-called
North Pacific. Yet Worm et al. find no strong evidence for this whaling/cascade hypothesis, Springer et al.’s (2003)
effect in the North Atlantic. In contrast to this last result, account for the North Pacific and southeastern Bering
Clapham and Link (Chapter 24) speculate that such rela- Sea, and a much earlier poorly known account by Barrat
tionships did occur in this area, thus suggesting that the lack and Mougin (1978) from the Southern Ocean. Branch and
of a significant finding in Worm et al.’s time series may to Williams (Chapter 20) build on these two reports and
some degree be caused by a temporal mismatch between the explore the feasibility for the Springer et al. hypothesis for
periods of whale removal (earlier) and the available time the Southern Ocean. Although evidence for causality is
series of fin fish and forage fish data. problematic (as emphasized by Ballance et al. in Chapter 17),
these authors conclude that the whaling/cascade hypothe-
Large Whales as Detritus sis is an easily feasible explanation for various southern
ocean pinniped population collapses (southern elephant
We turn next to the role of dead whales as detritus (Chapter 22). seals and southern sea lions in this case), and the hypoth-
Here Smith summarizes the empirical evidence for the esis is consistent with much of the available information.
significance of this process that has been obtained from Branch and Williams further conclude that recently
experimental and opportunistic studies of the coloniza- reported minke whale declines are unlikely to have been
tion, growth, and extinction of detritivores on dead whale driven by killer whale predation, although that conclu-
carcasses. The author emphasizes a remarkable possibil- sion is partly based upon the assumption that all or much
ity—that a single whale carcass may influence the nearby of the minke whale carcasses are consumed. The authors
seafloor community for decades. Smith also concludes that also point out that any explanation of these declines raises
this process probably is not of great significance to seafloor a large number of unanswered questions, and hence, like
ecosystems on continental shelves but that it may be the analyses by Doak et al. (Chapter 18), the contribution
extremely important in the deep sea. Another interesting by Branch and Williams is not so much an argument as it
dimension to this process is that, because of the great is a feasibility analysis. Chapter 21 by Mangel and Wolf
longevity of whale carcasses on the seafloor, deep-sea explores the Springer et al. hypothesis using optimal
ecosystems may just now be feeling the effects of the foraging theory and decision analysis, essentially asking
impoverishment of this resource that resulted from indus- the question: Does it make sense? These authors conclude
trial whaling. that the hypothesis is indeed plausible, but they also iden-
tify the need for additional information. Plausibility, of
Large Whales as Prey course, does not establish factuality. It does, however, in
the absence of empirical evidence, establish the ease or
The ecological role of large whales as prey is the most con- difficulty with which an ecological interaction might
troversial of the three potential food web pathways that we occur. This knowledge, in turn, provides a quantifiable
have identified. This is due in part to the nature of the evi- sense of likelihood as well as an indication of whether or
dence and in part to equally controversial indirect effects not the pursuit of additional empirical evidence is worth
of the interaction on other important components of the the effort.
food web (Springer et al. 2003). Several chapters of the vol-
ume bring a variety of interesting dimensions to questions The Broader View—Discipline, Timescale,
regarding the nature and magnitude of killer whale preda- Socioeconomics, and Process
tion on large whales. Reeves et al. (Chapter 14) summarize
the evidence and arguments for and against the view of The volume also includes a number of chapters that explore
killer whales as consequential predators of large whales, broader issues concerning whales and whaling in ocean
although they draw no firm conclusions. In Chapter 18, ecosystems from the perspectives of deeper time, other inter-
Doak et al. address the plausibility of the most controver- action web processes, policy, and law. Although this volume
sial aspect of Springer et al. (2003)—the question of the is about whales and whaling, not fisheries and oceanography,
nutritional importance of great whales to killer whales and it would be incorrect and unfair not to acknowledge the
how this changed as a result of whaling. Not surprisingly, influence of these disciplines on the way we look at ocean
the results depend strongly on the assumptions, but they ecology. Chapter 6, by Hunt, provides that perspective by dis-
illustrate that large whales could have provided a signifi- cussing the importance of bottom-up control, largely seen to
cant food resource for transient killer whales, even if attacks be driven by climate change and human exploitation. The
are rarely seen and if only limited tissues or segments of author gives a synopsis of the evidence for bottom-up forc-
large-whale populations were consumed. Readers interested ing effects and points out that these effects cannot be disre-
in the Springer et al. hypothesis are referred to recent garded in exploring the consequences of predator removals
and top-down effects. Further, Hunt notes that for marine

R E T R O S P E C T I O N A N D R E V I E W 391

mammals, the occurrence of top-down control appears to be tists may have placed too much emphasis on bottom-up
rare and that bottom-up and top-down forcing are processes forcing in their efforts to understand ocean ecosystem
whose importance in a given marine ecosystem is scale- dynamics. These authors also ask the most forward-looking
dependent. In Chapter 7, Lindberg and Pyenson explore the question in this volume: If the whales recover, what will
relationships between whales and ocean ecosystems in this mean to the oceans? Their conclusion is that it will
macroevolutionary time, from the Cretaceous-Tertiary (K-T) probably mean quite a lot.
boundary through the period of the fossil record of cetaceans
and their recent ancestors. These authors summarize a large Have We Achieved Our Goals in This Volume?
body of paleontological information, from which they
suggest that loss of marine reptiles at the K-T boundary had There is no simple answer to this question. A careful look
a strong influence on cetacean evolution, affecting such back through the chapters reveals that, in fact, there is little
fundamental characteristics as diet, body size, and species in the way of concrete empirical evidence for the ecologi-
interactions. This is a rich potential area for future research. cal effects of whales and whaling. There can be little doubt
Chapter 27 by Costa et al. would seemingly appear as an that the oceans have changed, but the degree to which
outlier to the volume, because it addresses pinnipeds rather whales and whaling caused or contributed to these
than whales or whaling. However, these authors present a changes is wrought with uncertainty and no small amount
global analysis of the patterns and reported causes of of controversy. It is difficult to separate the influences of
pinniped population declines and, in doing so, establish the whaling from the influences of oceanographic and climate
oceanic conditions facing the higher-trophic-level marine change, or from the effects of more recent or ongoing fish-
species, including whales. Costa et al. indicate that pinniped eries, especially given that so much attention has been
declines have occurred in various species and in many dif- paid to these latter influences and so little attention has
ferent areas. Despite many interesting patterns, especially been paid to the influences of whales and whaling. Viewed
relating to taxonomy, behavior, and physiology, the authors in this way, one might conclude that we have failed. But
also recognize perplexing inconsistencies. Thus, even under while the empirical evidence for the effects of whales and
the best of circumstances the actual causes of these declines whaling is thin for each case, a collective view of the
are in most cases poorly understood. combined empirical evidence, theory-based approaches,
and process-based evidence from other more tractable
Because whaling was a human endeavor with important species and ecosystems yields a substantially different inter-
social and economic dimensions, no volume on whales and pretation. This more synthetic approach leads to the
whaling would be complete without some attention paid to inescapable conclusion that whales have had large and
the question of what they meant and continue to mean to important effects on the structure, function, and evolution
people. The chapters by Bromley (Chapter 28) and Orbach (29) of ocean ecosystems and that whaling has altered the
discuss whaling from social, economic, legal, and policy per- structure and function of ocean ecosystems over the last
spectives. Two of us (D.P.D. and R.L.B.) note that the discus- two centuries. Viewed in this way, we believe that we have
sion of the IWC is more complex than described by Orbach succeeded. Although case-by-case arguments remain, no
when one factors in the issue of the Revised Management longer is it reasonable to think of whales simply as
Scheme, which is currently under development by the IWC. passengers in a changing ocean. The volume provides a
clear view of the processes by which whales and whaling
Finally, in Chapter 30 Kareiva et al. deal with the immediate can influence the oceans as well as a road map for how
problem of whale conservation, casting the main approaches, these processes can be further studied and understood.
findings, and arguments from various contributions in this
volume into the broader context of the discipline of ecology The Future
and the more general issues and needs of conservation. From
each perspective, the authors make the following important Considering what we know and don’t know about whales,
points. The first is that scientific culture has affected the whaling and ocean ecosystems, and looking now to the
manner in which we view the questions, and this very often future, what would we like to know and how should science
leads us somewhat astray. Second, while the problem of proceed? If nothing more, this volume highlights an impor-
whale conservation and recovery may seem dire, it is in fact tant point—that whales and whaling have influenced the
much more hopeful than similar issues on the land, where oceans in significant ways. As decisions are being made about
species are now extinct and habitats have been grossly how future whale stocks should be managed, this perspective
altered. Third, continued human exploitation is not the only must be added to the ever-evolving algorithms used to advise
threat to the recovery of whales. And lastly, part of what managers, if not brought to the fore. Although collectively
makes this entire problem interesting is the fact that whales we agree with Kareiva et al. (Chapter 30) that in the past there
are a human cultural phenomenon. Kareiva et al. go one step has been too much attention to numerology and not enough
further to chastise the cetacean research and conservation attention paid to ecology or, for that matter, other biological
community for placing too much emphasis on counting disciplines, individually we are of differing minds on the
and not enough emphasis on ecology, and they raise the
question of whether the larger community of ocean scien-

392 OVERVIEW AND SYNTHESIS

importance of these measures. Three of us ( J.A.E., D.F.D., the killer whale/marine mammal assemblages in different
and T.M.W.) believe that improved estimates of historical ocean basins are sustainable without historic abundance lev-
whale abundance and current carrying capacities will be nec- els of large whales, and whether population change is more
essary to properly inform management decisions because of sensitive to bottom-up or to top-down forcing processes. As
their central importance in the assessment of the potential pointed out by Doak et al., these approaches usually do not
ecosystem impacts of whaling and in the establishment of provide final definitive answers, but they do help establish
targets for recovery. Two of us (D.P.D. and R.L.B.) do not sup- the degree of plausibility of hypothesized processes and thus
port that view, instead believing that the central goal of are an important step in the quest for final answers.
research should be to obtain improved information on cur-
rent abundance, stock structure, and factors that limit the Finally, we encourage future cetologists to build their
growth rates of large whales and to distinguish between conceptual visions and conduct their research in the com-
changes in large-whale abundance caused by anthropogenic pany of interdisciplinary collaborators. There has been
impacts and those caused by environmental change. Achiev- some effort already to join forces with oceanographers, and
ing a capability for the implementation of either view will that of course should continue. But the dimensions to the
likely be problematic, but the beginnings of a path have been interplay between whales, whaling, and ocean ecosystems
established in many of the chapters found in this volume. are much bigger than this. We hope this volume has made
that point. We also note that whales are not so unique that
Looking beyond numerology, what about ecology, physi- our understanding of them cannot inform and be informed
ology , behavior, and evolution? What sorts of research might by comparisons with and insights from other taxa and the
be conducted in the future to understand better the influ- rest of ecology.
ences of whales and whaling on ocean ecosystems? One obvi-
ous approach is simply to watch ecosystem change as whale In conclusion, we note with interest the parallel paths
stocks recover, while keeping the confounding influences of Western culture has taken when attempting to place the roles
other factors such as fisheries and oceanographic change well of humans and whales in an ecological context. Both have
in mind. This is a potentially powerful research tool that can proven difficult, although perhaps for different reasons. It
be employed to test some of the current thinking that is both appears that it was easier for us to consider humans an impor-
hypothetical and controversial. As suggested by Clapham tant element of the marine ecosystem than it was for us to
and Link (Chapter 24), if we are to understand better the eco- consider large whales important. We hope this volume has
logical roles played by whales, future marine mammal sur- contributed to the position that whales and whaling are
veys must be integrated with multidisciplinary ecosystem aspects of the marine ecosystem that have to be integrated
research, notably in areas that are already well studied. They into the more traditional fields of fishery, wildlife biology,
further note that a parallel approach would be to conduct and oceanography. In the absence of such integration, many
such integrated surveys in simpler ecosystems where whales of the mysteries regarding how the world’s oceans function
have to date failed to recover from whaling, in order to mon- will remain enigmatic.
itor system changes as recovery occurs.
Literature Cited
A second approach to the problem is the increased use of
theory and interdisciplinary synthesis. Physiological limita- Barrat, A. and J.L. Mougin. 1978. The Southern elephant seal,
tions of the animals comprised in ecosystems represent a Mirounga leonina, of Possession Island, Crozet Archipelago,
powerful, yet sorely underutilized, avenue for predicting 46°25’ south, 51°45’ east. Mammalia 42: 143–174 (in French).
what can or cannot occur. Despite their size, whales can swim
only so fast when foraging and can process only so much DeMaster, D.P., A.W. Trites, P. Clapham, S. Mizroch, P. Wade,
food in a day when feeding. Such biological limitations and and R.J. Small. 2006. The sequential megafaunal collapse
traits define the limit of their influence and the resulting hypothesis: testing with existing data. Progress in Oceanography
strength of interactions between individual predators and 68: 329–342.
prey. Coupled with demographic and behavioral informa-
tion regarding prey choice, such interdisciplinary approaches IWC. 2005. Report of the Scientific Committee. Journal of
have changed how we view whales in ecosystems. The chap- Cetacean Research and Management (Supplement) 7: 32–34.
ters by Doak et al. (18), Williams (15), and Branch and
Williams (20) illustrate the power of this method, but they Springer, A.M., J.A. Estes, G.B. van Vliet, T.M. Williams,
are only a beginning. Modeling approaches could be used to D.F. Doak, E.M. Danner, K.A. Forney, and B. Pfister. 2003.
understand better such fundamental problems as whether Sequential megafaunal collapse in the North Pacific Ocean: an
ongoing legacy of industrial whaling? Proceedings of the
National Academy of Sciences 100: 12223–12228.

Whitehead, H. and R.R. Reeves. 2005. Killer whales and whaling:
the scavenging hypothesis. Biology Letters 1(4): 415–418.

R E T R O S P E C T I O N A N D R E V I E W 393



INDEX

NOTE: Page numbers followed by the letters f or t denote references contained in figures or tables.

aboriginal whaling style, 87 in maintaining biodiversity, 18–19 beluga whale, 82, 83
abundance overexploitation of, 10–11 Berger, Joel, 174
quantity vs. complexity of, 47 Bering Sea/Aleutian Islands, marine mam-
climate regime shifts and, 2 total catches of, 39f
declines in, xvi, 1 Archaeoceti, 68 mal biomass, 117–125
and ecological impact of whale decline, Arctic aboriginal whaling, 88–89 abundance estimates, 117–120
Arnould, John P. Y., 344 annual and seasonal biomass by sub-
27 Atlantic cod (Gadus mohua), 10, 53
estimates of, 2, 117–120 Australian fur seals, 350 group, 120–121
natural variations in, 11 composition, changes in, 122
albacore (Thunnus alalunga), 39 Baird’s beaked whale (Bernardius bairdii), 82 general patterns, 122–123
Altamira cavern, 29 baleen whales large whales, 121
American bison, 27, 28, 33, 380–381 management implications, 122–125
American-style pelagic whaling, 90–91 Bryde’s whale, 39, 41, 249f, 252f, 253 methods, 117–120
American-style shore whaling, 90 in Central North Pacific, 39, 41 parameters used in biomass calculations,
animal size, ecology of, 29. See also extreme historical population estimates of, 108t
humpback whale, 72, 83, 91, 103, 104, by species, 131–133t
body size percent reduction in, 120
Antarctic fur seal (Arctocephalus gazelle), 55, 107, 111–113, 121, 122, 134–135, population distribution, 119–120
174, 181–182, 248f, 251, 320 population declines, possible causes of,
350 molecular evolution of, 104
Antarctic minke whales, Southern Hemi- prey concentration necessary to support, 124–125
55 recommendations, 125
sphere, 266, 270–271, 272 sei whale, 39, 116, 142–143, 249f, 252f, results, 120–122
anthropogenic forcing, 11 253 seasonality, importance of, 122
anthropogenic influences whaling and, 82, 102 small cetaceans, 121–122
Ballance, Lisa T., 215 species included in, 118t
exotic species, introduction of, 20, 23, Barrett-Lennard, Lance G., 163 top predators, removal of, 116
68 Basque-style whaling, 87, 89–90 bigeye tuna (Thunnus obesus), 39
bay whaling era, 85 Bigg, Michael, 164
habitat fragmentation, 19–20, 68 bearded seal (Erignathus barbatus), 56, 122, biodiversity decline, 19, 23, 202
overhunting, 68 345 biomass dynamic models, limitations of,
apex predators
ability to change ecosystems, 20 45–46
depletion of by industrial fishing, 38 biomass pyramids, oceanic, 29–30
exploitation of, 47
and habitat fragmentation, 19–20

395

biophysical ecology, 373 changes associated with commercial crabeater seal (Lododon carcinophagus), 55,
blue whale (Balaenoptera musculus) whaling and fishing, 39, 42–48 56, 219

body size, 191 contemporary, 40–41 Croll, Donald A., 202
North Pacific Ocean, 248f, 251–252 historical, 41–42
as prey for killer whales, 174 species composition, 39–40 Danner, Eric M., 134, 245
bluefin tuna (Thunnus thunnus), 39 cetacaceans Darwin, Charles, 8
Bluhm, Bodil A., 303 body size, 74–75, 76f Davis, Lance, 365
body size, 29, 74–75, 76f, 109. See also composite phylogeny of, 69f DDT, 22
diversification patterns, 68, 69, 70–71, dead zones, 34
extreme body size dead-whale detritus
bottom-up control, 2, 8, 385, 391 73f
ecological role, 73 biodiversity and whale-fall species, 291,
defined, 67 evolution of, 68–75 293, 295t
Southern Ocean, 218–219 feeding styles, 73–74
sperm whales, 327–328 fishing up prey size, 76–77, 78 deep-sea effects of, 287–293
bottom-up forcing, 385 food webs, 69f enrichment-opportunist stage, 288, 289f,
bowhead whale (Balaena mysticetus) foraging and prey distribution, 55–56
evolution of, 55 morphology, 70f 290f
killer whale attacks on, 178–179, 183 oldest known, 68 as habitats for marine life, 383
killer whale phobia in, 179 physical factors influencing evolution of, in the intertidal, 294
migratory patterns of, 117 in marine ecosystems, 286–298, 391
North Pacific Ocean, 247f, 250, 254–255 72–73 mobile-scavenger stage, 288, 289f
population, 121, 122 population, bottom-up control of, 55–56 in the pelagic region, 294
whaling and, 83, 88–89, 91, 124 tectonics, 69f production and initial fate of, 287
Branch, Trevor A., 262 terrestrial-to-aquatic transition, 73 at shelf depths, 293–294
Bromley, Daniel W., 363 Cetotheriids, 71 sulfuphilic stage, 288, 289f, 290–291
Brownell, Robert L. Jr., 134, 215, 388 changing baseline perspective, 7 whale-fall succession in regions beyond
Bryde’s whale (Balaenoptera edeni), 39, 41, Channel Islands, trophic reorganization
California slope, 291
249f, 252f, 253 and, 20–22, 23 whaling, effects of on the production of,
Clapham, Phillip J., 174, 215
California sea lion, 351, 352f climate variability 294–298
Cape fur seals, 350 DeMaster, Douglas P., 116, 388
capture myopathy, 181 during evolution of cetacaceans, 72 density dependence, 385
capture stress, 181 and extinctions, 15 density manipulation, 8
carbon and fish populations, 52–53 detritus. See dead-whale detritus
in population control, 50 diet choice model, 279–281
global cycling of, 8, 9 responses to, 58 D-loop, genetic divergence in, 104–105
sinking of to sea floor, 34 co-evolutionary model, 363–365 Doak, Daniel F., 231, 388
Caribbean monk seal (Monacus tropicalis), apologetics, 365, 372 Donlan, C. Josh, 14
elaboration and reconstitution, 364–365 drift whales, 87
22 naturalization, 365 dwarf minke whale (Balaenoptera acutoros-
carousel feeding, 167 resource, 363–364
carrying capacity, 32–33, 55, 103, 111, 338, commercial whaling, 82, 83. See also indus- trata subspecies), 191

374, 382 trial whaling; whaling ecology, hypothesis testing and falsification
cascade, general conditions of, 285 food web structure changes associated in, 33
case studies
with, 42–48 EcoSim modeling technique, 11, 41
ecosystem effects of fishing and whaling future of, 368–371 ecosystem(s). See marine ecosystems; ocean
in the North Pacific and Atlantic large-whale community composition,
Oceans, 335–342 ecosystems
changes to, 207, 209 Ecotrophic Efficiency in Ecopath model, 41
gray whales in the Bering and Chuckchi recovery and, 369–370 Edenic Ideal, concept of, 371–372
Seas, 303–311 rise and fall of, 365–368 eel grass (Zostera marina), wasting disease of,
suspension of, 368
potential influences of whaling on pin- sustainable resumption of, 370–371 9
niped populations, 344–356 consequential interactions, in large, open effective size, estimates of, 109
El Niño-Southern Oscillation (ENSO)
sperm whales in ocean ecosystems, ecosystems, 9–11
324–333 press perturbations, 10–11 events, 2, 53–54
pulse perturbations, 9–10 endangered list, removal from, 102
whales, whaling, and ecosystems in the conservation, 374, 392 Endangered Species Act (ESA) of 1973, 371,
North Atlantic Ocean, 314–322 contributors, list of, vii–viii
controlled manipulation, 8 374
Caspian seals, 348 coral reef ecosystems ENSO events. See El Niño-Southern Oscilla-
catastrophic mortality events, ecological Caribbean web, 31
decay of, 22 tion (ENSO) events
consequences of, 10 northwestern Hawaiian Islands (NWHI) environmental movement, growth of, 374
catch histories era, defined, 85
and main Hawaiian Islands (MHI) Eskimo bowhead whaling, 88–89
attempts to determine, 92–94 compared, 30–31 Essington, Timothy E., 38
intentional underreporting, 94 overfishing and, 22 Estes, James A., 1, 231, 388
primary sources, 93 sponges and, 33 Exclusive Economic Zone, establishment of,
secondary sources, 93 Costa, Daniel P., 344
catch records, accuracy of, 113 Coyle, Kenneth O., 303 371
Central North Pacific (CNP) exotic species, introduction of, 20, 23, 68
defined, 39 exploitation
food web structure
apex predators, 47
biomass dynamic model to study, as a benefit, 28
39–40

396 I N D E X

industrial fishing as, 38 water temperature and, 52, 53 carrying capacities past and present, 32–33
whaling as, 38, 42, 82, 134 zooplankton abundance and, 52, 53 historic and modern nesting beaches,
extinction food web(s)
and apex predators, loss of, 18–19 changes to associated with commercial map of, 32f
Atlantic gray whale, 317–318 greenhouse gas emissions, 34
causes of, determining, 14 whaling and fishing, 39, 42–47 Greenpeace, 380
climate change, role of, 15 community-level structuring forces in, 67 grizzly bear (Ursus arctos), predation by,
ecosystem simplification in, 18 competition, 47
human-driven, 15, 16, 24, 27–28 defined, 7 175–176
keystone herbivore hypothesis of, 16 destabilization of, 34 groundfish, 52–53, 336
limpet, 9 disruption of due to megafauna declines,
mass events, 28 habitat fragmentation, 19–20, 68
megafaunal, 16 28 harbor seal (Phoca vitulina richardii)
overkill hypothesis of, 15, 16 fishing down, 10, 22, 23f, 76
passenger pigeons, 33 human-induced changes in, 383 killer whales and, 262
Pleistocene, 15–18 pathways, 2, 8 population trends, 348, 349f
extreme body size. See also blue whale (Bal- removal of large whales from, conse- whaling and, 116
harp seal (Pagophilus groenlandica)
aenoptera musculus); killer whale (Orci- quences of, 134, 210–211 food limitations of, 56
nus orca) reorganization, implications of, 14 population, 55, 345
and ability to maintain metabolic func- structure of in Central North Pacific slaughter of, 31
harvesting quotas, 134
tions, 192 (CNP), 40–42 hawksbill turtle, 33
advantages of, 192 whales and, xv–xvi, 2, 69f, 383 Heise, Kathy A., 163
caloric demands and, 193–194, 197–198 forage fish, 336 Hewitt, Roger P., 215
constraints on, 193–196 Forney, Karin A., 145 Highsmith, Raymond C., 303
digestive pathway, 195 freshwater ecosystems, studies of, 8–9 historical ecology, 28
fueling in the wild, 196–199 functional webs, 7 hooded seals, 345
heart mass, 194 fur seals human(s)
hunters vs. grazers, 192–193 killer whales and, 262 civilization, growth of, xv
individual organs, size of, 195–196 and removal of large whales from the and decline of megafauna, 27–28, 29
intestinal length and body length, 195 and pinniped depletion, 355
physiological and ecological conse- Southern Ocean, 223–224 as structuring agents in ecosystems, 38,
Southern Hemisphere, 264–265
quences of, 191–200 355–356
terrestrial animals with, 192 Gallman, Robert, 365 values, reconciliation with public policy,
genetic divergence rate, 104–105
factory ship whaling, 92 genetic diversity 375–376
false killer whale (Pseudorca crassidens), 181 humpback whale (Megaptera novaeangliae)
fecundity, 51, 52 estimates of, 108–109
figures, list of, xi–xiv in fin whales, 107 estimated population of, 103
fin whale (Balaenoptera physalus) in humpback whales, 107 feeding sites, 72
maintenance of, 134 genetic diversity in, 107
carrying capacity, 55 management implications related to, genetic studies of, 104
genetic diversity in, 107 and killer whales, 174, 181, 182
genetic studies of, 104 111 migratory patterns of, 134
heart size, 194 in minke whales, 107 North Atlantic Ocean, 320
killing of, concern for, 245 at silent positions of protein-coding North Pacific Ocean, 248f, 251
North Pacific Ocean, 141–142, 249f, population estimates, 111, 112f, 113,
genes, 106
252–253, 255 genetic drift, 103, 104 116, 121, 122
population estimates, 103, 111, 113, 122 genetic sweeps, 104 spatial population structure, 134–135
spatial components of harvest and Gilmore, Raymond M. 178 whaling and, 83, 91
Gleiter, Karin, 365 Hunt, George L. Jr., 50
decline of, 141–142, 143 gray whale (Eschrichtius robustus) hybrid food web modeling approach, 11
whaling and, 83 hypermutation, 106–107
fisheries Atlantic population, extinction of,
ecological impact of, 8–9, 10 317–318 industrial fishing
sustainability of, 28 and depletion of apex predators, 38
fishing distribution/abundance of in Bering and ecological impact of, 10–11
and depletion of apex predators, 38 Chukchi seas, 303–304, 309–311 exploitation by, 38
effects of on marine ecosystems,
feeding industrial whaling. See also commercial
335–341 biological and physical impacts of, 309 whaling; whaling
strategies, evaluation of, 11 primary foods of, 306–309
fishing up prey size, 76–77, 78 sites in the Bering and Chuckchi seas, depletion of large-cetacean populations
fish populations 72, 303–306 by, 262–264
bottom-up effects on, 52–54
climate and, 52–53 killer whale attacks on, 177–178, ecosystem and populations, effect on,
depletion of, 10 181–182, 183 262
El Niño-Southern Oscillation (ENSO)
in North Pacific Ocean, 250–251 hypothetical effects of on marine ecosys-
events, 53–54 population decline, 121, 122, 311 tem, 340f
and relative mobility of predator and removal of from endangered list, 102
“snorkeling” by, 178 killer whale predation habits, effect on,
prey, 57 spyhopping, 178 262–263
whaling and, 88, 91, 303
Greater Yellowstone Ecosystem, predator in the North Pacific, 22
pinnipeds and cetaceans in Aleutian
eradication, effects of, 19
green turtle (Chelonia mydas), 22 archipelago prior to, 198f–199f

INDEX 397

industrial whaling. See also commercial coexistence of with possible prey, 184 resistance to attack, 181–182
whaling; whaling (continued) consumption patterns and limitations, sea otters, 2, 10, 22, 23, 196, 231–232,

population spatial structure, study of, 182–183, 239–240 233
135–143 description of, from prey’s perspective, selection, determinants of, 198–199
size/age preferences in, 181
catch, 136, 137 163 sperm whales, 328–329
catch-per-unit-effort (CPUE), 135, diet and foraging behavior, 77–78, Steller sea lions, 231–232, 233
prey switching, 169–170
136, 137–138, 141f 167–170, 174, 263, 286t, 269–272 pygmy, 181
Directed Effort Window (DEW), 136 distribution/abundance of, 145–158, 164 as reason for megafaunal collapse, 279
effort, 136–137 resident-type, 146, 147, 163, 164, 165f,
fin whales, 141–142 Indian Ocean, 151
implications for management of, North Atlantic and adjacent seas, 166, 167, 168, 169, 174, 182
role of tradition in lives of, 170
139–143 154–155 scavenging behavior, 177
main components of, 135 North Pacific Ocean and adjacent Arc- seasonal movement patterns, 145
methods, 135–137 social structure of, 166–167
results, 137–139 tic waters, 150–151 sperm whale effects on, 331–332
sei whales, 142–143 patterns of, overview, 150, 157–158 subdivision of ecotypes into regional
spatial grid, 135–136, 139f South Atlantic, 155, 157
species studied, 135 South Pacific, 151 populations, 164–166
species harvested, biomass density, Southern Hemisphere, 266–267 surplus killing, 169
Southern Ocean, 157 sympatric ecotypes in Northeastern
140f worldwide density, 156f
intact terrestrial ecosystems, study of, 29 fish predation, 167–168 Pacific Ocean, 164
interaction webs, 7, 8–9 heart size, 194 time to consume biomass of prey by
interdisciplinary synthesis, need for, 393 intentional beaching by, 168
International Convention for the Regula- interaction of with other marine mam- species, 197f
top-down control by, 56, 67
tion of Whaling, 367–368 mals, 174 transient-type, 146, 147, 163, 164,
International Whaling Commission (IWC), K-T impact event, 77–78
large whales in diets of, 183–184 166–167, 168, 169, 174, 178, 182,
103 mammal-eating, 146 (see also transient 391
annual catch records, 135 vocalizing by, 180
creation of, 367 killer whales) whaling and, 82, 226
harvest data, 246 marine mammal declines in relation to Komodo dragon (Varanus komodoensis), 18
life history parameters, 327 Konar, Brenda, 303
management procedures, 111 dietary needs, Southern Hemi- Krill Surplus Hypothesis, 217–218
moratorium on whaling, 375, 382 sphere, 274 Kudela, Raphael, 202
recommendations re: pre-whaling abun- marine mammal predation by, 168–169,
174–175, 262, 274, 353, 355 La Brea tar pits, 16
dance estimates, 118, 124 metabolic needs of, 238 Lago Guri reservoir, impact of, 18–19, 23
0.54K benchmark, 382 metabolism, 271 lance-and-wait technique, 88
interspecific interactions, 20, 21f natural history and ecology of, 163–171 large whales
in North Pacific Ocean, 256–257
Jackson, Jeremy B. C., 27 oceanic and neritic, 147 Bering Sea/Aleutian Islands, 121
offshore-type, 146, 147, 165f, 167 current populations, trophic importance
Kalland, Arne, 380 phobia, in bowhead whales, 179
Kareiva, Peter, 379 population estimates, 146, 147–150 of, 210
Kauffman, Matthew J., 134 line-transect survey (LTR), 148t, 149 daily metabolic rates, 204t, 205, 209–210
kelp forests, overgrazing of, 1, 22 mark-recapture (MR) method, 148t, decline of, community implications,
keystone herbivore hypothesis of extinc-
149 210–211
tion, 16 non-standardized survey (SURV), 148t, as detritus, 391
keystone species effects, 10 ecological impact of removing from the
killer whale (Orcinus orca), 51, 116 149–150
observation and anecdotal informa- North Pacific, 202–212
Antarctic Type C, 147 individual whale prey biomass con-
approximate ranges of, by type, 165f, tion (OBS), 148t, 150
photo-identification catalog (CAT) sumption, 205
166f net primary production, 205
attacks method, 147, 148t, 149 primary production required to sustain
by region, 152–153t
behavior during, 180–181 population structure, 163–166 whole populations, 205–206,
on bowhead whales, 178–179 position of in food chain, 174 208t, 209t
firsthand descriptions of, 175–176 prey rapid removal, 207, 209
on gray whales, 177–178, 181–182, alternative, Southern Hemisphere, results, 205–207
trophic level and trophic transfer effi-
183 273–274 ciencies, 205, 207t
rates, 238–239 Antarctic minke whales, Southern whale population estimates, 203, 205
scarring and mutilation as evidence whale prey biomass consumption,
Hemisphere, 270–271, 272 203, 204t, 205
of, 182 anti-predation strategies by, 170 ecological impact of removing from the
on sperm whales, 179–180 large baleen and sperm whales, Southern Ocean, 215–228
surge in reports of, 184 commercial whaling by species, 217t
behavioral versatility of, 170 174–184 conclusions, 227–228
body size, 75 large cetaceans, Southern Hemisphere, fur seals, 223–224
caloric demands of, 197–199, 200 history of area whaling, 217
coastal fish-eating, 146–147 (see also resi- 273
large whales, skepticism concerning,
dent killer whales)
183
pinnipeds, Southern Hemisphere, 270,

271–272

398 I N D E X

ice seals, 219–220 coupling diet choice and prey popula- muted cascade, 10
Krill Surplus Hypothesis, 217–218 tion dynamics, 281 Myers, Ransom A., 335
loss of prey for killer whales, 226
overview of, 215 in North Pacific, predation and, narwhals, whaling of, 82, 83
penguins, 221–226 232–234 natural resource, defined, 364
pinnipeds and seabirds, 220–221, neotropical anachronisms, 1
parameter estimates and sample calcu- net whaling, 88
223–224 lations, 281–283 non-standardized survey (SURV), 148t,
post-whaling trends, 218–226
Ross Sea Shelf and Slope areas, predictions, 283–284 149–150
in Southern Hemisphere, 274–275 North Atlantic Oscillation (NAO) atmos-
226–227 ecological roles of, 116
as predators, 390–391 evaluating impacts of commercial harvest pheric circulation pattern, 53
as prey, 391 North Atlantic right whale (Eubalaena
Lascaux cavern, 29 on, 123–124
law and culture, relationship between, heart mass in relation to body mass, glacialis), 90, 182, 318–320
North Pacific gray whale (Eschrichtius robus-
373–374 194f
less-than-zero-sum ecology, 34 international attitudes toward, 375 tus), 55
limpet (Lottia alveus), extinction of, 9 intestinal length in relation to body North Pacific Ocean
Lindberg, David R., 67
line-transect survey (LTR), 148t, 149 length, 195f blue whales, 248f, 251–252
longline catch rates, decline of, 47 killer whale predation by, 168–169 bowhead whales, 247f, 250, 254–255
Lotze, Heike K., 335 metabolic rate in relation to body mass, Bryde’s whales, 249f, 252f, 253
commercial whaling, effects of,
MacArthur, Robert, 9 193f
Magnuson, Senator Warner, 371 migratory patterns, 117 257–258
Magnuson-Stevens Fisher Conservation Act most trophically distinct, 326t depletion of whales in, 245–246, 248
public interest in, 379–380 fin whales, 249f, 252–253, 255
(FCMA) of 1976, 374 sequential collapse of in North Pacific gray whales, 250–251
Mangel, Marc, 279 history of whaling in, 246–249
marine bird populations Ocean, 263f
mark-recapture (MR) method, population nineteenth century, 246
at-sea distribution, 55 postwar whaling, 248
bottom-up control of, 54–55 estimates, 148t, 149 twentieth century prior to World
and El Niño-Southern Oscillation Martin, Paul S., 14
mass extinctions, 28 War II, 247–248
(ENSO) events, 54 maximum sustainable yield (MSY), princi- humpback whales, 248f, 251
and food limitation, 54 killer whales, 256–257
foraging and prey availability, 55 ple of, 374 marine mammals, sequential collapse of,
penguins, 221–226 megafauna
and removal of large whales from the 263f
in Africa, 29 minke whales, 253
Southern Ocean, 220–221 climate changes and decline of, 28 right whales, 247f, 249–250
marine ecology defined, 27 sei whales, 249f, 252f, 253
ecological consequences of the loss of, 28 sperm whales, 250f, 253–254, 255–256
absence of large animals, implications humans and the decline of, 16, 27–28, 29 whale depletion in, 258
of, 30 megafaunal collapse, and killer whale pre- whales harvested within 100 nautical

baseline community concept, 30 dation, 279 miles of the coast, 257f
basic principles, origins of, 29 Melville, Herman, 324 North Pacific right whale (Eubalaena japon-
biological oceanographers vs. ecologists mesonychids, 68
microbial loop, 30, 34 ica), 88, 124
and fisheries biologists, 30 MIGRATE program, 108 northern bottlenose whale (Hyperoodon
biomass pyramids in, 29–30 migration
marine ecosystems. See also ocean ecosys- ampullatus), 82, 92
age- or sex-specific patterns of, 117 northern elephant seal, 348
tems estimates of, 108 northern fur seal (Callorhinus ursinus), 116,
anthropogenic influences on, 19–20, 23, seasonal variations in, 117
to avoid predation by killer whales, 170 122, 350
68 minke whale (Balaenoptera acutorostrata) Norwegian-style shore whaling, 84, 91–92
coral reef, 22, 30–31 Antarctic, 266, 270–271, 272 no take rule, 374, 375
effects of fishing and whaling on, 1, carrying capacity, 55 nutritional limitation hypothesis, 2
dwarf, 191
296–298, 335–341 genetic diversity in, 107 observation and anecdotal information
freshwater, 8–9 migratory patterns of, 135 (OBS), 148t, 150
history of whale populations as back- North Pacific Ocean, 253
population estimates, 111 ocean ecosystems. See also marine ecosys-
drop for management of, population structure of, 135 tems
102–103 whaling and, 88
human alterations, overview of, monk seals, 33, 34 biodiversity loss and, 202
335–336. See also fishing; whaling Muir, John, 374 bottom-up effects on, 2
management, 376 multispecies relationships. See interaction climate change and, 51
methodical dismantling of, 33 El Niño-Southern Oscillation (ENSO)
Marine Mammal Protection Act (MMPA) of webs
1972, 374–375, 374 mutation rate events, 2
marine mammals. See also pinnipeds evolution and history of, 2
bottom-up effects on, 55–56 extreme rate heterogeneity, 106 fisheries removal and, 51
decline of genetic divergence, 104–105 importance of whales to, 382–384
classic diet choice model as explana- hypermutation, 106–107 large predators in, 196–199
measuring, 104–107 management of, 376
tion for, 279–281, 284 variation, cautions concerning, 105–106 removal of marine mammals from,

effects of, 116

INDEX 399

ocean ecosystems. See also marine ecosys- southern hemisphere species, 349–350 Reeves, Randall R., 82, 174
tems (continued) temperate and tropical species, resident killer whales, 146, 147, 163, 164,

role of sperm whales in, 211–212, 348–349 165f, 166, 167, 168, 169, 174, 182
324–333 and removal of large whales from the resource, defined, 363–364
right whale (Eubalaena glacialis)
social systems and, 363–372 Southern Ocean, 220–221
co-evolutionary elaboration, model sensitivity of to environmental fluctua- North Pacific Ocean, 247f, 249–250
for, 363–365 population estimates, 111
future of commercial whaling, tions, 344 pygmy, 191
368–371 time spent foraging at sea, 354f ringed seal (Pusa hispida), 122, 192, 345
recovery, 369–370 whaling, effects of, 344–356 Roemer, Gary W., 14
rise and fall of commercial whaling, worldwide distribution of, 344–345 Roman, Joe, 102
365–368 pirate whaling operations, 92 Ross Sea Shelf and Slope, 226–227
Pitman, Robert L., 215
whales and whaling, effects on, xv, 2–3, Pleistocene extinctions Scammon, Charles, 163, 174
7, 8, 27 bottom-up vs. top-down views of, 17 sea grass, disappearance of, 9
causes of, 15 sea lions, 350–351
oceans ghosts of predators past, 18 sea otter (Enhydra lutris)
history of, xv implications of for present-day ecology,
lack of collective public wisdom and Bering Sea, 116
common sense about, 28 15–18 decline of, 1–2, 22, 125
in oceans, 22–23 influence of on kelp forest ecosystems, 1,
O’Connor, Casey, 376 predator-prey relationships, 16
offshore killer whales, 146, 147, 165f, 167 poison whaling, 87–88 10, 22
omnivory, 31 polar bear (Ursus maritimus), 116, 192 killer whales and, 231–232, 233, 262
Omura, Hideo, 245 polychlorinated biphenyls (PCBs), 381 overexploitation of, 10
open-boat whaling, 85, 329 population control as prey for killer whales, 168, 196
operation, defined, 85 availability of prey, 58 as sea urchin predators, 1, 57
optimum sustainable population (OSP), bottom-up mechanisms for, 50, 51, sea urchins (Diadema antillarum)
catastrophic collapse of, 9–10
principle of, 374 52–57, 58 as prey for sea otters, 1, 57
Orbach, Michael K., 373 climate variability and, 50, 58 seabirds. See marine bird populations
overexploitation, 14 disease as, 51 sei whale (Balaenoptera borealis)
growth rate, 52 Bering Sea, 116
apex predators, 10–11 lack of safe refuges, 57, 58 Central North Pacific, 39
Atlantic cod, 10 predation and, 50–51 North Pacific Ocean, 142–143, 253
cetaceans, 56 and relative mobility of predator and spatial components of harvest and
sea otters, 10
overfishing, 1, 11 prey, 57, 58 decline of, 142–143
community-wide impacts of, 31 resource availability and, 50, 51 selective species culling, negative conse-
ecological effects of, 22–23 scale dependence of bottom-up and top-
extreme global, 31 quences of, 31–32
overharvesting, x, 22, 202, 350 down mechanisms, 56–57 sensitivity analyses, 113
overhunting, 68 top-down mechanisms for, 50–51, 57, 58 sharks, 10–11
overkill hypothesis of extinction, 15, 16 population monitoring, 134 shoot-and-salvage approach to whaling, 87
population spatial structure, 134 short-finned pilot whale (Globicephala
Paine, Robert T., 7 predator control. See top-down control
Palumbi, Stephen R., 102 predator-prey disequilibria, 19 macrorhynchus), 180, 181
passenger pigeons, 33, 380 predators short-term local harvesting, effects of, 134
Peche Merle cavern, 28–29 eradication of, 19 shrimp (Pandalus borealis), depletion of, 10
penguins, and removal of large whales turning into prey, 20–22 Siniff, Donnald B., 215
preservation, 374 skipjack tuna (Katsuwanus pelamis), 39, 42
from the Southern Ocean, 221–226 press perturbations small-type whaling, 92
Pfister, Beth, 116 deep-water, benthic systems (Atlantic small-world networks, 28, 33
photo-identification catalog (CAT) method, Smith, Craig R., 286
cod-shrimp interactions), 10 Smith, Tim D., 82
population estimate, 147, 148t, 149 oceanic, totally pelagic systems (sharks, South American sea lion, 351
Piatt, John F., 245 southern elephant seals, 265–266,
pilot whale, 75, 82, 180, 181 tuna), 10–11 Southern Hemisphere
Pinchot, Gifford, 374 shallow-water, benthic systems (sea
pinnipeds fur seals in, 264–265
otters), 10 killer whales in, 263, 266–267, 268t,
bottom-up control of, 55–56 prey switching, 169–170
decline of, 2 production control. See bottom-up control 269–275
dive performance, 354f public policy rise and fall of megafaunal populations
overharvesting of, 350
population declines and human values, reconciliation with, in, 264–266
375–376 Southern Ocean
human intervention and, 355
life history and behavioral correlates, whaling and, 371–372, 374–375 commercial whaling in, history of, 217
“pull of the recent,” 72 defined, 215, 216f
351–353 pulse perturbations Ross Sea Shelf and Slope areas, 226–227
potential causes of, 351–355 top-down vs. bottom-up control,
risk of predation, 353, 355 Diadema dieback, 9–10
population trends sea grass disappearance, 9 218–219
Arctic species, 345, 348 Pyenson, Nicholas D., 67 southern sea lions, Southern Hemisphere,
otariid species, 350–351 pygmy killer whale (Feresa attenuata), 181
overview of, 345, 346t–347t, 355–356 pygmy right whale (Caperea marginata), 191 265
pygmy sperm whale, 326 species preference law, 375

400 I N D E X

sperm whale (Physeter macrocephalus) Trivelpiece, Wayne Z., 215 spatial structure of, 134
Bering Sea, 116 trophic cascade, 8, 10, 19, 31, 210 synthesizing historical views of, 111–113
Central North Pacific, 39 trophic level, 51, 56 whale prey biomass consumption
competitors, 326–327, 331, 333 trophic reorganization, 20–22 estimates of, 203, 204t, 206t
demands of on marine ecosystems, trophic structure, energy flow and, 50 individual, 205, 209–210
211–212 tropical aboriginal whaling, 89 whale population, 205, 210
density-dependent responses in, tuna, 10–11 whales. See also individual species by
327–328, 333 turtlegrass, 33
estimated number caught annually, 330f name
evolution of, 324–325, 333 United States Endangered Species list, 102 abundance vs. environmental fluctua-
feeding habits of, 41, 325–326 upper-trophic-level organisms, bottom-up
and killer whales, 174, 179–180, tions, 385–386
331–332 limitations on, 56 beaching of, 382
life history, 327 upper-trophic-level predator, defined, 51 comeback of selected species, implica-
migratory patterns of, 117
miscellaneous ecological effects of, 332 Van Vliet, Gus B., 245 tions of, 384–385
natural predation on, 328–329 as detritus, 2, 3, 286–298
North Pacific Ocean, 250f, 253–254, Wade, Paul, 145 ecological roles, lack of information
255–256 Walker, Theodore J., 178
in ocean ecosystems, 324–333 walrus (Odobenus rosmarus divergens), 116, about, 382, 392, 393
population future of, 392–393
bottom-up control, 327–328, 332 117, 122 history of, need to know, 102
estimates for, 121, 122, 211 water temperature, and fish populations, iconic state of in conservation lore,
human regulation of, 330
top-down control, 329 52, 53 379–380
predation by, 211–212, 330–331 Weddell seals (Leptonychotes weddellii), 220 importance of
pygmy, 326 Weise, Michael J., 344
rosette formation, 179, 180 whale carcasses. See dead-whale detritus to ocean ecosystem functioning,
trophic value, 42 whale ecology, xv 382–384
whale falls, 297f whale population(s)
whaling, effects of, 82, 91, 93, 124, weight of evidence argument for, 11
329–330 back-calculation models, requirements interest in, reasons for, 388–389
for, 125 of the North Pacific
sponges, 33
spotted hyena (Crocuta crocuta), predation baleen whales, historical estimates of, current and pre-exploitation estimates
108t of, 234t
by, 176
Springer, Alan M., 245 benchmarks to evaluate recovery, 102 demographic models for, 235–236
spyhopping, 178 declines in, possible causes of, 124–125 modeling dynamics and deaths of,
starvation, 52, 54 demise of, spatio-temporal variation in, 3
Steller sea cow (Hydrodamalis gigas), 22, 116 ecological impacts of changes in, 236–237, 238t, 240
Steller sea lion (Eumetopius jubatus) numbers and life histories of, 234–235
123–124 as sufficient food for killer whales,
Bering Sea, 116 estimates, 203, 205
decline of, 124–125, 196–197, 351 238–242
indirect fishing, effects on, 384 bias in, 125 as just another overharvested natural
killer whales and, 231–232, 233, 262 effective size, 109
population estimates, 122 historic and global, 383t resource, 380–381
stranded whales, 87 hunting mortality in, 113 as predators, 254–256
subsistence whaling, 82, 83, 314 killer whales, 147–150 as prey (top-down control), 2, 3
super-whale, 380 line-transect survey (LTR), 148t, 149 as prey for other predators (bottom-up
supply-side ecology, 385 mark-recapture (MR) methods, 148t,
surplus killing, 169 control), 2, 3, 256–257 (see also
149 killer whales)
tables, list of, ix–x migration of, 108 recovery
take, defined, 374 non-standardized survey (SURV), 148t, constraints on, 381–382
temperate aboriginal whaling, 89 lessons to be learned from, 386
Terborgh, John, 19 149–150 role of in an ecosystem context, 254–257
terrestrial carnivores, predatory behavior observation and anecdotal informa- whaling. See also commercial whaling;
industrial whaling
of, 175–176 tion (OBS), 148t, 150 aboriginal, 87
Tershy, Bernie R., 202 photo-identification catalog (CAT) American-style pelagic era, 90–91
Tinbergen, Niko, 279 American-style shore era, 90
top predator, defined, 51 method, 147, 148t, 149 ancient activities, lack of information
top-down control, 2, 8, 11, 385, 391–392 pre- vs. post-whaling, 111–113 about, 92
sensitivity analyses in, 113 annual catches in the Southern Hemi-
defined, 67 underestimation, 124 sphere and northern North Pacific,
Southern Ocean, 218–219 genetic (DNA) approaches to, 103–104, 295f
sperm whales, 329 Arctic aboriginal era, 88–89
transient killer whales, 146, 147, 163, 164, 113 baleen whales, 82, 102
genetic data and analysis, reliability of, Basque-style era, 89–90
166–167 168, 169, 174, 178, 182, 391 bowhead whales, 83, 88–89, 91, 124
tritrophic food chains (TFCs), 31 109–111 catch histories, 92–94
genetic divergence rate, 104–105 as a cause of megafaunal collapse, 2
genetic diversity in, 103–104, 108–109, data, sources of, 94f
defined, 82
111 ecosystem effects of, xv, 1, 2–3, 7, 8, 27,
maximum sustainable yield, 103 296–298, 314–322, 335–341, 389
prey biomass needed to support, 191 efficiency in, 84
projected, for 2050, 381t equipment, method, and techniques, 84
questions regarding, 389–390
recovery of, 3, 103

INDEX 401

whaling. See also commercial whaling; locations of, 85f shoot-and-salvage approach, 87
industrial whaling (continued) minke whale, 88 small-type era, 92
moratoriums on, 102, 375, 382 in the Southern Ocean, 217
eras, 85–87 narwhals, 82, 83 species involved in, 82
approximate time periods, 88f net era, 88 sperm whales, 82, 91, 93, 124, 329–330
summary list of, 86–87f in the North Atlantic Ocean, 314–322 subsistence, 82, 83, 314
in the North Pacific Ocean, 246–249, sustainable harvest, 375
Eskimos’ equipment and techniques, 87 temperate aboriginal era, 89
ethnic/national groups involved in, 83 257–258 tropical aboriginal era, 89
as exploitation, 38, 42, 82, 134 North Pacific right whale, 124 and whale populations, depletion of, 38
factory ship era, 92 northern bottlenose whale, 82 Whitehead, Heal, 324
fin whale, 83 Norwegian-style shore, 84, 91–92 wildfires, 28
future of, 392–393 operations, 83–84, 85f Williams, Terrie M., 191, 231, 262, 388
geographic locations of, 83 operations, list of grouped by era, Wolf, Nicholas, 279
gray whale, 88, 91, 303 wolverine (Gulo gulo), predation by, 176
Greenpeace interference with, 380 99–101f Worm, Boris, 335
history of, 82 pelagic/shelf/intertidal effects of,
humpback shale, 83, 91 yellowfin tuna (Thunnus albacares), 39
International Convention for the Regu- 295–296 Younger Dryas (YD) cold snap, 15
pinnipeds and, 116, 344–356 Yuan-Farrell, Christopher, 376
lation of, 367–368 pirate operations, 92
International Whaling Commission poison era, 87–88 zero-sum ecology, 8, 9, 11, 33–34
in production of whale detritus, 294–295 zooplankton abundance, and fish popula-
(IWC), 103, 111, 118, 124, 135, public policy with respect to, 371–372,
246, 327, 367, 375, 382 tions, 52, 53
killer whale, 82, 226 374–375
lance-and-wait technique, 88 reasons for, 83–84
law and culture, 373–376 seasonality and years of, 84

402 I N D E X