Download Place-Based Ecosystem Management in the Open Ocean

MCBi.qxd
2/2/2005
2:41 PM
18
Page 302
Place-Based Ecosystem Management in
the Open Ocean
Elliott A. Norse, Larry B. Crowder, Kristina Gjerde, David
Hyrenbach, Callum Roberts, Carl Safina, and Michael E. Soulé
Rancher, about the Texas landscape, to former ship’s captain: “Did you ever see
anything so big?”
Former ship’s captain: “Well, yes. A couple of oceans.”
Texas rancher: “Well I declare! Oceans. Hmmmph!”
William Wyler’s 1958 film, The Big Country
Our modern human brain evolved in the African sa-
using hand-nets, buckets, and scuba, oceanogra-
vanna, where our ability to learn the rewards and per-
phers and fisheries biologists have relied mainly
ils of its vast landscape was the key to our survival.
on remote sampling technologies, which have led
Now that humans have expanded far beyond our
them to focus on aggregated metrics of ecosystem
motherland, our success depends on our ability to
function (e.g., plankton productivity) and popu-
handle the rewards and perils of ecosystems very dif-
lation structure (e.g., fish biomass), rather than
ferent from the one that shaped us. Of these, none is
the intricacies of behavioral interactions within
larger, more complex, nor more difficult for people to
and among species.
fathom than the blue and black waters of the open
ocean.
Our terrestrial species pays little attention to the
open ocean for several reasons:
1. It is remote from human observers, out of sight,
hence out of mind.
2. To our unattuned eyes, its wavy surface seems impenetrable, featureless, and trackless.
3. Conducting research there is much more expensive and requires more labor, expensive equip-
5. Large pelagic animals are uncommon in the open
ocean and often move quickly, so are seldom seen
alive and are far less known by scientists, decision
makers, and the public than nearshore species.
6. The seeming scarcity of humans in the vastness
of oceanic ecosystems makes the open ocean
seem invulnerable to human impacts.
7. Compared with estuaries and enclosed seas,
the open ocean shows much less impact from
proximity to dense human populations.
ment, and logistical coordination than doing re-
8. Sixty-four percent of the sea (and a much larger
search in intertidal, estuarine, and inshore areas.
percentage of the open ocean beyond the conti-
4. In contrast to marine biologists’ in situ research
302
nental margins) lies on the high seas, outside of
MCBi.qxd
2/2/2005
2:41 PM
Page 303
Place-Based Ecosystem Management in the Open Ocean
303
the potential protection conferred within
Sargassum, corals, squids, sea turtles, seabirds, pinni-
nations’ exclusive economic zones (EEZs).
peds, and whales are not fishes.
Yet the open ocean is far from biologically homogeneous (Boehlert and Genin 1987; Polovina et al.
Effects of Fishing in the Open Ocean
2000; Worm et al. 2003), and there is pressing need to
In recent years, scientists have published unambigu-
modify the way we manage this vast ecosystem be-
ous evidence that fishing has depleted populations
cause its megafauna—its equivalents of tigers, bears,
and disrupted ecosystems in coastal waters worldwide
wolves, eagles, condors, crocodiles, rhinos, and ele-
( Jackson et al. 2001). The evidence is no less com-
phants—are vanishing, while its most vulnerable ben-
pelling for the open ocean. For instance, North At-
thic ecosystems are increasingly being degraded. A di-
lantic humpback (Megaptera novaeangliae), fin (Bal-
verse, growing chorus of informed voices (e.g., FAO
aenoptera physalus), and minke (B. acutorostrata) whale
2003; Gislason et. al 2000 and papers that follow; Juda
populations appear to be 85 to 95 percent below for-
2003; Mooney 1999; Pew Oceans Commission 2003;
mer population levels (Roman and Palumbi 2003).
Pikitch et al. 2004; Ward and Hegerl 2003) explain why
Biomass of fishes has declined precipitously since 1900
ecosystem-based management is needed to stop the
in the North Atlantic (Christensen et al. 2003). The
loss of biodiversity and the collapse of fisheries or offer
trophic level of fish catches has declined progressively
ways to implement it. This chapter discusses place-
since the 1950s (Pauly et al. 1998), at the same time
based methods of ecosystem-based management—in-
that fishing has pushed ever deeper (Pauly et al. 2003).
cluding fishery closures, marine protected areas, and,
Populations of oceanic tunas and billfishes appear to
in particular, marine reserves—for protecting and re-
have decreased about 90 percent worldwide since the
covering biological diversity in the open ocean, and
1950s (Myers and Worm 2003). North Atlantic oceanic
specifically, their application to two different kinds of
sharks have decreased more than 50 percent in the last
ecosystems, seamounts and the epipelagic zone.
8 to 15 years (Baum et al. 2003); oceanic whitetip
Humans can reduce populations of oceanic species
sharks (Carcharhinus longimanus) (Figure 18.1) in the
and alter oceanic ecosystems via pollution (oil and
Gulf of Mexico have decreased more than 99 percent
other toxic chemicals, nutrients, noise, solid wastes),
since industrial longlining began there in the 1950s
ship strikes, climate change, and, possibly, introduc-
(Baum and Myers 2004). Moreover, while the world’s
tion of alien species (e.g., toxic phytoplankton, jelly-
fishing power has dramatically increased, the world’s
fishes). There is reason for concern about activities
marine fish catch — corrected for overreporting by
that could occur in coming decades (e.g., deep-sea
China, the largest fishing nation—has actually de-
mining, deliberate ecosystem manipulation via iron
creased since 1988 (Watson and Pauly 2001). The
fertilization, deep-sea disposal of carbon dioxide,
United Nations’ latest global assessment (FAO 2002)
ocean thermal energy conversion). But the most per-
considers 75 percent of the world’s marine capture
vasive and severe human impacts in the open ocean
fisheries to be fully exploited or producing less than
for which there is compelling scientific documenta-
they would at lower fishing levels. Fishing is emptying
tion is the killing of targeted pelagic and benthic spe-
the oceans of its large predators.
cies and its consequences for other species and habi-
These declines reflect a profound shift in predator–
tats. For lack of a precise term applicable both to
prey dynamics, the predator in this case being hu-
species targeted by fisheries and associated “collateral
mankind. Oceanic fisheries have undergone dramatic
damage” to nontarget species and habitats, we use the
technological advances in recent decades, as fishermen
term fishing throughout this chapter, recognizing that
have adopted roller and rockhopper trawls (which
MCBi.qxd
2/2/2005
304
2:41 PM
Page 304
Marine Conservation Biology
F I G U R E 1 8 . 1 . Oceanic whitetip shark (Carcharhinus longimanus). A recent study (Baum
and Myers 2004) found that more than 99 percent of these once-ubiquitous predators
have been eliminated from the Gulf of Mexico since the advent of industrialized
oceanic longlining in the 1950s. This ecological extinction of large predators can
start trophic cascades in marine ecosystems, as it can in freshwaters and on land.
Photo © Masa Ushioda, CoolWaterPhoto.com.
gave them access to previously unfishable de facto
significant as functioning components of their eco-
refuges, including the steep, rocky slopes of sea-
systems. Fishes, in contrast, have not become signifi-
mounts), nearly invisible drift gillnets tens of kilome-
cantly more capable of avoiding capture.
ters long, and pelagic longlines of comparable length
Of course, biological extinction is worse still, because,
that couple bait with fish-attracting light-sticks, all of
among other reasons, it precludes any possibility of re-
these now made of strong, nondegradable synthetic
covering from commercial or ecological extinction.
materials. Other powerful fishing technologies include
Fishing has caused the biological extinction or en-
steel hulls, big engines, precision depth finders and
dangerment of large marine species (Carlton et al.
fish finders, detailed mapping of the seafloor, floating
1999; Dulvy et al. 2003; Roberts and Hawkins 1999;
fish-aggregating devices with radio beacons used by
Myers and Ottensmeyer, Chapter 5) and threatens
purse seiners, electronic net sensors, real-time down-
many more. For instance, concerns about severely re-
loads of satellite-derived sea surface temperature im-
duced white marlin (Tetrapturus albidus) and northern
agery showing the locations of fronts, global position-
bluefin tuna (Thunnus thynnus) populations have
ing systems, and at-sea processing. Armed with this
prompted proposals for their listing as endangered
suite of technologies, people have become ever more
species (Rieser et al., Chapter 21). The prospects for
capable of fishing species to commercial and ecologi-
seamount species could be even worse, given their as-
cal extinction (Myers and Ottensmeyer, Chapter 5), ef-
sociation with fixed habitats, their vulnerable life his-
fectively making them uneconomical to pursue and in-
tories, their high endemism, and the rapid spread of
MCBi.qxd
2/2/2005
2:41 PM
Page 305
Place-Based Ecosystem Management in the Open Ocean
305
trawling into the deep sea. However, lacking sufficient
the local extirpation of their prey (Crooks and Soulé
knowledge about the status of many taxa, it is unlikely
1999), and (3) the local disappearance of many smaller
that scientists could satisfy restrictive legal definitions
animals (prey) by competitive exclusion (Henke and
of endangerment for seamount species before they
Bryant 1999).
were severely depleted. Indeed, because they are
Terborgh et al. (1999) describe trophic cascade ef-
highly aggregated, serial depletion of fish populations
fects on new islands in Lago Guri, Venezuela, in which
seamount by seamount — each initially yielding a
predators disappeared and many kinds of herbivores
high catch per unit of effort (CPUE)—could easily
became superabundant, increasing herbivory on
mask overall abundance trends until all these habitat
seeds, seedlings, and leaves and causing complete re-
patches are overexploited. The insensitivity of the
cruitment failure for most tree species. The extirpation
CPUE statistics to the actual population abundance
of wolves (Canis lupus) from Yellowstone National
for patchily distributed species requires precautionary
Park (USA) in the 1920s led to a severe increase in
management, including the establishment of marine
browsing pressure by elk (Cervus canadensis) on young
protected areas (MPAs).
quaking aspen (Populus tremuloides) and willow (Salix
In addition to the harm caused by overfishing of
spp.), the primary food of beaver (Castor canadensis).
targeted populations, bycatch and entanglement in
Aspen failed to recruit into the canopy for 80 years or
fishing gear threaten many oceanic species. Some of
so, causing the entire beaver wetland ecosystem to dis-
these, including leatherback sea turtles (Dermochelys
appear from Yellowstone’s northern range (Ripple and
coriacea), short-tailed albatross (Phoebastria albatrus),
Larsen 2000).
and North Atlantic right whales (Eubalaena glacialis)
Neither evidence nor theory indicates that trophic
are at high risk of extinction and are officially pro-
cascades are confined to the land. In coastal eco-
tected under the US Endangered Species Act and the
systems, Jackson et al. (2001) show that elimination of
Convention on International Trade in Endangered
key species catalyzes equally dramatic ecosystem
Species of Wild Fauna and Flora, and are on the World
transformations. There is no reason to believe that re-
Conservation Union (IUCN) Red List. For too many
moval of large predators would have less significant
species, however, such designations have not arrested
ecosystem-level consequences in the open ocean, and
their slide toward extinction.
there is intriguing reasoning suggesting that removal
Moreover, the loss of a species is not only conse-
of species has significant top-down effects (Verity et al.
quential in an evolutionary sense; the reduction or
2002), although these are more difficult to detect.
elimination of its ecological interactions can lead to
Knowing that serial overfishing has systematically re-
the disappearance of many other species (Soulé et al.
moved high trophic level species worldwide (Pauly et
2003). As on land, two groups of strong interactors in
al. 1998), the first question that should be posed is,
marine ecosystems that are particularly important to
How does that removal affect ecosystem function-
conserve are ones that have large effects on food webs
ing? Unfortunately, the two sciences that have most
and those that provide habitat structure. Terrestrial
influenced marine resource management have largely
ecologists have shown that removal of large predators
failed to address these questions. Biological oceanog-
has profound effects on the key ecosystem attributes of
raphy has focused mainly on bottom-up studies of nu-
species composition, structure, and functioning. The
trient and plankton dynamics, while fisheries biology
cascading consequences of species removal include: (1)
has focused mainly on single-species stock-recruit-
overbrowsing and subsequent changes in vegetation
ment modeling of commercially exploited species.
structure and composition (Soulé and Noss 1998), (2)
These sciences have largely overlooked top-down ef-
the ecological release of middle-size predators causing
fects resulting from deletion of megafauna, which,
MCBi.qxd
2/2/2005
306
2:41 PM
Page 306
Marine Conservation Biology
along with habitat disturbance, is the most important
concern about the effects of trawling recently
global marine ecosystem change currently under way.
prompted the release of an unprecedented statement
This oversight is ironic because scientific understand-
of concern by 1,136 marine scientists and conserva-
ing of the keystone role played by certain predators
tion biologists from 69 countries (MCBI 2004).
arose from studies in marine systems that were pub-
Given the incontestable evidence that fishing
lished prominently more than three decades ago
sharply reduces oceanic biomass, populations of large
(Paine 1966, 1969). Yet, a recent check of Science Cita-
species, average trophic level, and benthic structural
tion Index Expanded on April 4, 2004, showed that,
complexity, unless a century of accumulated ecologi-
among the 500 most recent (1998–2004) scientific pa-
cal theory and empirical evidence is wrong, there is no
pers citing Paine’s classic 1966 paper, only 5 are in
longer any credible denial that fishing profoundly al-
fisheries journals (and several of those papers concern
ters oceanic ecosystem functioning.
freshwater ecosystems).
The challenge of oceanic conservation requires ac-
In fact, the keystone role played by large predators
knowledging the impacts of fishing on open ocean
in North Pacific kelp forest ecosystems was demon-
ecosystems. Even where nations manage fisheries
strated by ecologists as early as the 1970s (Dayton
within their EEZs, the stock assessment-based para-
1975; Estes and Palmisano 1974). Since then, Estes et
digm, which focuses on estimating individual species’
al. (1998) and Springer et al. (2003) have elucidated
fishing mortality and controlling it by using catch
additional trophic links between nearshore kelp
quotas, size limits, gear restrictions, and temporary
forests and oceanic ecosystems. Ecological theory and
area closures, has produced few successes (two un-
a growing body of evidence indicate that removal of
usual ones are discussed by Hilborn, Chapter 15). A
oceanic megafauna has induced a serial trophic cas-
substantial fraction of oceanic fisheries occur outside
cade that has ramified throughout a vast area of the
of nations’ jurisdictions. International fisheries regu-
North Pacific for decades. It is difficult to explain
latory organizations are even less likely than those of
how such momentous findings have had so little ef-
individual nations to be effective in conserving target
fect on marine policy and management.
species and biodiversity due to competing national in-
In recent years it has also become clear that bot-
terests and the global “tragedy of the commons.” Fur-
tom trawling is much like forest clearcutting, in that
thermore, ensuring compliance in the oceanic realm
it removes the ecosystem’s structure-forming species
is even more difficult than doing so in coastal waters,
(Watling and Norse 1998; Watling, Chapter 12). Their
with enforcement on the high seas so uncommon that
removal has profound effects on benthic ecosystems
the pursuit and capture of a Uruguayan vessel, the
(Auster and Langton 1999; Dayton et al. 1995; Steele
Viarsa (Fickling 2003), one of the many vessels ille-
2002). Loss of biogenic structure eliminates habitat
gally fishing for Patagonian toothfish (Dissostichus
for species — including the young or adults of some
eleginoides), made global news.
commercially important fishes (e.g., Auster et al.
To summarize, open ocean ecosystems are severely
2003) — that associate with seafloor structures to
imperiled despite nations’ and international organi-
avoid predation, feed, or breed. Trawling on sea-
zations’ management efforts to date. We believe there
mounts off Tasmania has apparently eliminated 95
needs to be a different management approach, a shift
percent of structure-forming species including corals
toward ecosystem-based management built upon the
and sponges (Koslow et al. 2001). Structure-forming
growing scientific understanding of oceanography
animals living in food-poor, cold, deep waters of con-
and the interconnections among populations in
tinental slopes, midocean ridges, and seamounts are
oceanic ecosystems, an approach that makes full use
probably especially slow to recover from severe dis-
of new scientific understanding and new technolo-
turbance (Watling and Norse 1998). The growing
gies. Perhaps the simplest, most robust tool for ecosys-
MCBi.qxd
2/2/2005
2:41 PM
Page 307
Place-Based Ecosystem Management in the Open Ocean
307
tem-based management is the place-based approach,
place-based approach for maintaining and recovering
protecting certain places temporarily or permanently
oceanic species and their ecosystems. Although our
from some or all preventable harm. In this chapter we
findings should be relevant to many oceanic eco-
define reserves as places that are permanently pro-
systems, we focus primarily on two contrasting types,
tected from all preventable anthropogenic threats.
seamounts and the epipelagic (upper 100 m of the
We use the qualifying term preventable here because
water column) zone, although fishing pressure on
place-based approaches cannot eliminate all anthro-
continental slopes, on midocean ridges, and in the
pogenic threats and impacts; some pollutants, alien
mesopelagic zone (100–1000 m depth) has also in-
species, pathogens, and effects of global warming do
creased in recent decades (Merrett and Haedrich
not respect the boundaries people draw (Allison et al.
1997). So far, fishery closures and MPAs that bar cer-
1998; Kim et al., Chapter 9; Lipcius et al., Chapter 19).
tain activities (e.g., oil and gas operations) are far
Reserves are the most protected of many kinds of
more prevalent than fully protected reserves in the
MPAs, which we define as areas permanently pro-
sea. Roberts and Hawkins (2000) estimate that less
tected against at least one preventable threat. Fishery
than one-hundredth of 1 percent of the sea is fully
closures, another place-based management tool, differ
protected in marine reserves. And because most are
by having narrower aims rather than broader biodi-
coastal and very few are truly oceanic (indeed, there
versity and ecosystem integrity goals, and are gener-
are no fully protected reserves in international waters),
ally temporary. They are often used for rebuilding
assessing the potential benefits of oceanic reserves is
overfished target populations or for protecting vul-
inherently difficult. Thus we must winnow insights
nerable life history stages of target species until fish-
from temporary fishery closures in the oceanic realm,
ing can be resumed. Reserves, other kinds of MPAs,
from MPAs in the open ocean that allow some kinds
and fishery closures are all place-based tools that can
of fishing, from fully protected marine reserves in
be used in ocean zoning (Norse, Chapter 25). We be-
shelf waters, and from protected areas in nonmarine
lieve that networks of protected places that are con-
realms.
nected through currents and movements of organisms
can become effective, self-sustaining tools for avoiding biodiversity loss.
Features in a Featureless Ocean
In recent years a fast-growing body of theory and
On seeing the wavy sea surface stretching to the hori-
evidence has indicated that marine reserves can be a
zon, it is easy to think of the oceans as uniform, a view
powerful ecosystem-based management tool for pro-
subliminally reinforced by the featurelessness of
tecting and restoring biodiversity and fisheries
oceans on most maps (Ray 1988). Yet, one of ocean-
(Conover et al. 2000 and papers that follow; Gell and
ography’s most important contributions to marine
Roberts 2003; Goñi et al. 2000 and papers that follow;
conservation biology has been documenting the
Houde 2001; Lubchenco et al. 2003 and papers that
ocean’s remarkable heterogeneity. Some of this het-
follow; Murray et al. 1999; Norse 2003 and papers that
erogeneity is driven by topographic features that affect
follow; but also see Lipcius et al., Chapter 19). These
the overlying water column, but many other patterns
studies, however, generally address intertidal or
arise from horizontal and vertical water movements
nearshore ecosystems. Whether their conclusions can
(Boehlert and Genin 1987; Haury et al. 2000; Roden
be extrapolated to oceanic ecosystems needs to be de-
1987). Topographic and oceanographic heterogeneity
termined. Indeed, there is probably no more stringent
create a mosaic of oceanic biodiversity, one different
test for the efficacy of the place-based approach than
from, but comparable to, terrestrial mosaics. In turn,
its application in the open ocean.
many oceanic species concentrate at ecotones, which
In this chapter we examine the potential of the
allow them to use different sets of environmental
MCBi.qxd
2/2/2005
308
2:41 PM
Page 308
Marine Conservation Biology
conditions that homogeneous ecosystems do not
their sessile, sedentary, and resident inhabitants. As a
offer.
result, seamounts are often colonized by structurally
complex communities of suspension-feeding sponges,
Seamount Oceanography, Biology
and Fisheries
corals, crinoids, and ascidians (Rogers 1994) that are
Seamounts are islandlike biological hot spots in the
tend to be dominated by deposit-feeders.
rare or absent from surrounding abyssal plains, which
open sea, in part because they provide a rare resource:
Seamounts are also biomass hot spots because they
hard substrate. The hypsographic curve (http://www
alter the surrounding water flow in ways that en-
.seafriends.org.nz/oceano/ocean13m.gif) shows that
hance local productivity (Boehlert and Genin 1987;
53.5 percent of the Earth’s surface (75.3 percent of the
Roden 1987). Submerged seamounts and oceanic is-
seafloor) is composed of ocean depths between 3 and
lands (seamounts that break the surface) often in-
6 kilometers. These are mainly abyssal plains covered
duce leeward plumes of elevated productivity due to
with fine muds (Seibold and Berger 1993). But inter-
localized upwelling, a phenomenon called the island
rupting the abyssal seafloor are active or extinct vol-
wake effect. Even in the absence of upwelling, sea-
canoes, sometimes isolated, sometimes in chains or
mounts might support high animal biomass because,
clusters. Tens of thousands of these seamounts rise a
like shallow fringing coral reefs, they offer a combi-
kilometer or more toward shallower, sunlit waters
nation of strong currents and structurally complex
(Rogers 1994), many of them breaking the surface to
seafloor habitat. This allows resident fishes both to
form oceanic islands. The tallest, Mauna Kea on
feed on passing zooplankters and small fishes (e.g.,
Hawaii (USA), which is, indeed, the tallest mountain
lanternfishes, Myctophidae) in the water column and
on Earth, rises nearly 10 km from its abyssal plain. Be-
to take refuge amidst seamount structures when ma-
cause vertical gradients and stratification create
rauding pelagic sharks or tunas arrive. The fishes and
pelagic depth zones with different conditions for life,
other animals that feed on passing organisms and de-
seamounts, which transcend these zones, create their
tritus effectively increase seamounts’ filtering area,
own benthic depth zonation.
providing more food to benthic communities than
The ecosystems on steep seamount flanks and pin-
might otherwise occur.
nacles are different from those continental shelves
Seamount surveys have found extremely high (>30
and slopes at similar depths, in part because waters
percent) levels of apparent endemism (Richer de
that bathe seamounts have much lower sediment
Forges et al. 2000), perhaps due to distinctive sea-
loads, allowing photosynthesis to occur at greater
mount circulation phenomena called Taylor columns.
depths. Indeed, the depth record for benthic algae,
These large stationary eddies increase the residence
268 m, is from a seamount (Littler et al. 1985). Water
time of overlying waters. Taylor columns of sufficient
clarity over seamounts must also affect the success of
duration would allow eggs and larvae to stay within
visual predators that feed on vertically migrating zoo-
waters above a seamount until individuals can settle.
plankton (Genin et al. 1994; Haury et al. 1995).
Having essentially closed populations with negligible
No less important, because seamounts can inter-
gene flow would favor seamount animals that be-
cept ocean currents, causing 100 to 10,000 times more
come reproductively isolated endemics. Among the
mixing than in waters farther from them (Lueck and
first adaptations that might be selected are shortened
Mudge 1997), turbulence resuspends and removes
larval duration or behaviors that would retain larvae
fine sediments from seamount slopes and pinnacles.
within Taylor columns.
On these sediment-free outcroppings, organisms can
The combination of topographically induced
settle and thrive without risk of clogging and burial.
oceanographic phenomena and hard substrates makes
Currents also deliver food to and remove wastes from
seamounts islandlike oases in desertlike oligotrophic
MCBi.qxd
2/2/2005
2:41 PM
Page 309
Place-Based Ecosystem Management in the Open Ocean
309
oceans. Not surprisingly, the abundance of demersal
were even less resilient to fishing than coastal species.
seamount life and distinctive oceanographic phe-
Orange roughy, a long-lived (maximum age >100
nomena attract highly migratory pelagic predators
years, maturity at 22–40 years) fish associated with
including cetaceans, seabirds, sharks, tunas, and bill-
seamounts and banks, exemplifies the inefficacy of
fishes. Seamounts also serve as rendezvous where
today’s fisheries management for conserving oceanic
some epipelagic (e.g., scalloped hammerhead sharks,
species (Branch 2001; Koslow et al. 2000). The story is
Sphyrna lewini; Klimley 1995) and deep-sea (e.g., or-
essentially the same for precious corals, spiny lobsters,
ange roughy, Hoplostethus atlanticus; Bull et al. 2001)
and pelagic armourhead: fishermen discover a large
fishes from wider areas converge to mate or spawn.
concentration, quickly deplete it, and move on, while
Of course, in a world driven by economics, no
the population either recovers very slowly or fails to
concentration of useful life goes unnoticed forever.
recover (Roberts 2002). Fisheries in deep-sea eco-
Benthic and demersal seamount species that fisheries
systems are unlikely to be sustainable for target species
target include pink and red precious gorgonian corals
or biodiversity in general (Merrett and Haedrich 1997;
(Corallium spp.), black corals (Antipathidae), gold corals
Roberts 2002). But the fact that seamounts are fixed,
(Gerardia spp.), spiny lobsters ( Jasus spp.), orange
discrete features and are demonstrably being seriously
roughy, hoki (Macruronus novaezelandiae), oreos (Ore-
impacted makes them the “low-hanging fruit” of
osomatidae), pelagic armourhead (Pseudopentaceros
oceanic conservation. Temporary fishery closures are
wheeleri), rockfishes, wreakfish and hapuka (Polyprion
unlikely to be effective in maintaining their biodiver-
spp.), and Patagonian toothfish. Most of these are
sity because their benthic communities are resident
long-lived, late-maturing species, traits that make
and long-lived. Protecting them in no-trawling MPAs
them especially vulnerable to overexploitation
or managing them as fully protected reserves are at-
(Roberts 2002; Heppell et al., Chapter 13). Indeed, Pa-
tractive alternatives to existing management tools
cific rockfishes and gold corals can reach ages up to
(Probert 1999).
200 years (Cailliet et al. 2001) and 1,800 years (Drufbottom longlines, but most often trawls—the fishing
Epipelagic Oceanography, Biology,
and Fisheries
gears that cause the most collateral damage to seafloor
The vast expanses of the water column also have bio-
communities (Chuenpagdee et al. 2003)—are used.
logical hot spots; some pelagic areas have far higher
Because habitat-forming corals and sponges are both
species diversity (Worm et al. 2003) or productivity
vulnerable to trawl damage and long-lived, recovery
(Polovina et al. 2000) than others. Phenomena that
from trawling can take years, decades, or centuries,
cause heterogeneity away from topographic highs in-
much like recovery from forest clearcutting (Watling
clude upwelling at divergences, fronts such as con-
and Norse 1998; Watling, Chapter 12). This makes
vergence zones between water masses and eddies spun
trawling the greatest threat to the world’s seamount
off from ocean currents (Hyrenbach et al. 2000). Up-
benthic and demersal species. The pelagic fishes that
welling brings nutrients from deeper waters into the
are caught over seamounts are the same species we ex-
euphotic zone. This stimulates phytoplankton pro-
amine in the next section.
duction above background values, leading, in turn, to
fel et al. 1995), respectively. Some fishes are taken with
The development of industrialized fishing in the
zooplankton blooms, which attract planktivorous
decades after World War II allowed fishermen to re-
squids and fishes such as anchovies (Engraulidae),
duce fish populations in coastal waters, compelling
lanternfishes, small carangids and their predators such
them to journey further and deeper to find new “un-
as common dolphins (Delphinus delphis), Cory’s shear-
derutilized” species to replace ones they had depleted.
waters (Calonectris diomedea), Atlantic blue marlin
Unfortunately, the new targets of distant water fleets
(Makaira nigricans), bigeye thresher sharks (Alopias su-
2/2/2005
310
2:41 PM
Page 310
Marine Conservation Biology
perciliosus), and Humboldt squid (Dosidicus gigas). Var-
a.
ious frontal features between water masses with different temperatures or salinities concentrate oceanic
predators and their prey (Figure 18.2). Among three
possible reasons are (1) steep temperature or water
clarity gradients are barriers to movement, (2) convergence flow at fronts aggregates buoyant or weakly
swimming prey, or (3) temperature gradients allow
epipelagic species to feed in food-rich cooler waters
and then accelerate digestion and growth in warmer
waters. Convergence zones between water masses are
often visible as surface drift lines. Detached seaweeds,
logs, and flotsam collect at these fronts, providing
b.
Survey effort (time)
Flying birds
Feeding / sitting birds
hard substrates for invertebrates such as Lepas gooseneck barnacles and attracting small fishes, young
loggerhead sea turtles (Caretta caretta), and larger predators such as dolphinfish (Coryphaena hippurus),
yellowfin tunas (Thunnus albacares), blue (Prionace
glauca) and oceanic whitetip sharks. Frontal features
may be essentially permanent, seasonally predictable,
or unpredictably episodic. Even those that are permanent can move hundreds of kilometers throughout
the year or from year to year (Hyrenbach et al. 2000).
Large oceanic species, including blue whales (Balaenoptera musculus), northern elephant seals (Mirounga
anguirostris), Laysan albatrosses (Diomedea immutabilis), leatherback and loggerhead sea turtles, albacore tuna (Thunnus alalunga), bluefin tunas, and mako
sharks (Isurus spp.), appear to divide their lives between loitering (and apparently feeding) in places
with elevated chlorophyll concentrations (Block et al.
2002; Hyrenbach et al. 2002; Polovina et al. 2000) and
moving quickly across expanses of oligotrophic waters, where they apparently feed very little. Food-rich
patches are often transitory, disappearing when discontinuities break down or when predators deplete
prey abundance. Many large oceanic species spawn,
nest, or calve only in certain places, perhaps where
they can optimize the balance between food availability and predation risk for their young. To reach
breeding areas they journey hundreds or even thousands of kilometers.
Fisheries target pelagic species including Pacific
Proportion (%)
MCBi.qxd
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
Onshore On-front Front Off-front Offshore
Location
F I G U R E 1 8 . 2 . Aggregation at fronts by short-tailed
shearwaters (Puffinus tenuirostris). 18.2a: Approximately
9 to 20 million short-tailed shearwaters migrate from
their breeding grounds in Tasmania (Australia) to the
Bering Sea, where they aggregate along tidal fronts to
feed on euphausiid swarms during the boreal summer
(photo: D. Hyrenbach). 18.2b: North of St. Paul Island,
Alaska (USA) in August 1989, 78 percent of short-tailed
shearwaters feeding or sitting on the water were sighted
along a 7.2 km frontal zone, with far fewer birds sighted
along 3.6 km zones immediately inshore and offshore.
Aggregations of feeding shearwaters within this narrow
frontal zone regularly surpassed 1,000 birds km2, with
dense flocks reaching densities up to 10,000 to 25,000
birds km2. Data from Hunt et al. (1996).
MCBi.qxd
2/2/2005
2:41 PM
Page 311
Place-Based Ecosystem Management in the Open Ocean
311
salmon (Oncorhynchus spp.), Atlantic salmon (Salmo
by Sladek Nowlis and Friedlander, Chapter 17. Ac-
salar), tunas, wahoo (Acanthocybium solandri), and
cording to Botsford et al. (2003), “For species with
swordfish (Xiphias gladius). Oceanic drift gillnetting
high rates of juvenile and adult movement, individ-
and longlining operations also kill many animals with
uals spend too much time outside of reserves for the
little or no market value, including marine mammals,
reserves to provide sufficient protection.” These sci-
albatrosses, petrels, and sea turtles. Although some
entists’ views are echoed by some managers; for in-
large oceanic species (e.g., dolphinfish) are fast-grow-
stance, the US Gulf of Mexico Fishery Management
ing and early maturing, which makes them relatively
Council (1999) states, “Marine reserves are most ap-
resistant to overfishing, many are slow-maturing and
propriate for species that are relatively sedentary, such
long-lived (Musick 1999; Wooller et al. 1992; Heppell
as snappers and groupers, or for species that have spe-
et al., Chapter 13), making them, like demographi-
cific nursery sites, such as coastal sharks. Reserves are
cally similar demersal seamount species, particularly
likely less appropriate for migratory species, such as
vulnerable to overfishing.
mackerel, tuna and billfish.”
Protecting, recovering, and sustainably exploiting
In recent years, however, scientists and policy ex-
epipelagic species is an extraordinary challenge be-
perts have begun to consider marine reserves in the
cause these species are wide ranging, their habitats
open oceans (e.g., Gjerde and Breide 2003; Hooker
move, they live far from observers and enforcers, and
and Gerber 2004; Hyrenbach et al. 2000; Mills and
fishermen have an ever-growing arsenal of powerful
Carlton 1998). Moreover, resource management agen-
technologies to catch them. Moreover, as with sea-
cies such as the US National Marine Fisheries Service
mount species, the majority of the realm where they
have been compelled to close large areas to US pelagic
dwell—the high seas—is the least protected part of
fishing off Hawaii, the South Atlantic states, and New
the Earth’s surface.
England (Department of Commerce 1999, 2000, 2001)
to protect nontarget organisms such as billfishes and
The Need for Place-Based Conservation in
the Open Ocean
sea turtles. A modeling study of Mediterranean hake
(Merluccius merluccius) led Apostolaki et al. (2002) to
conclude that “yield and SSB [spawning stock bio-
The concept of protecting places has fundamentally
mass] benefits can be obtained through the use of a
changed conservation worldwide on land, in fresh
marine reserve even for highly mobile fish and un-
waters and, increasingly, in nearshore waters. But
derexploited fisheries.” The US National Research
until recently there has been little evidence that sci-
Council (Houde 2001) finds that, “When the mobility
entists view protecting ecosystems as a viable conser-
of adults is high, as in many pelagic and migratory
vation tool for the open ocean. For example, Angel
fish species, reserves have often been discounted as an
(1993) says, “The scales of oceanic systems are so
effective management tool . . . [but] even for highly
large that the methodologies developed for terrestrial
migratory species such as swordfish (Xiphias gladius)
conservation are inapplicable.” Agardy (1997) be-
or tunas, MPAs that protect nursery areas or vulnera-
lieves that “it is notoriously difficult to attach bound-
ble population bottlenecks may be effective manage-
ary conditions to marine ecological processes . . . this
ment tools.”
holds true not only for open ocean pelagic environments but for coastal zones as well.” The idea of
epipelagic reserves is dismissed by some experts. Par-
Challenges to the Place-Based Approach
rish (1999) believes that for species with highly
In response to the loss of biodiversity and resources
pelagic or migratory behavior, marine reserves will do
that humans value economically, place-based conser-
little toward achieving optimum yield, a view shared
vation began to extend from the land into nearshore
MCBi.qxd
2/2/2005
312
2:41 PM
Page 312
Marine Conservation Biology
marine ecosystems in the 20th century. Increasing
migrate, forage, and breed, reserves could protect
recognition that the entire world ocean is losing many
habitat hot spots by incorporating a novel design
of its large animals means that humankind will need
concept: dynamic boundaries.
to extend protection of places into the open ocean in
Let us examine the diversity of considerations in es-
the 21st century. The greatest challenge to place-based
tablishing networks of protected places in the open
conservation in the open ocean, as with conservation
oceans.
in general, is not so much scientific understanding as
the human dimensions, especially political commit-
Highly Migratory Species
ment: Do people who make key decisions consider life
A central question is whether ecosystem-based man-
on Earth important enough to protect and recover it
agement can help conserve highly migratory species
in the face of pressure to continue current practices?
that visit places only sporadically. The fact that large
For the open ocean the portents are mixed, ranging
pelagic species concentrate in certain places at certain
from somewhat encouraging (International Whaling
times—which makes them fishable in waters where
Commission, Convention on the Conservation of
their average density is very low—also allows their
Antarctic Marine Living Resources) to woefully disap-
protection in appropriately designed reserves, other
pointing (International Commission for the Conser-
MPAs, or fishery closures. Roberts and Sargant (2002)
vation of Atlantic Tunas).
suggest five ways in which migratory species can ben-
Even if legislators and managers decide that biodi-
efit from fixed-boundary marine reserves. (1) Many
versity loss and collapsing fisheries must be avoided,
migratory species travel through physical bottlenecks
there are significant scientific, technological, eco-
during their life cycle—for example, when they ag-
nomic, and legal challenges to place-based conserva-
gregate to spawn—and fisheries often target them in
tion in the open ocean. Because most of the open
these vulnerable locations. Reserves, less protective
ocean lies in international waters, questions concern-
MPAs, or temporary fishery closures sited in these
ing responsibility for protecting oceanic biodiversity
bottlenecks could significantly reduce fishing mortal-
are likely to loom larger than scientific and techno-
ity. (2) Single-species closed areas have already been
logical challenges. Strong international agreements
shown to boost yields of migratory species through
and mechanisms to ensure compliance are needed for
protection of juveniles from premature capture; for
place-based conservation to become an effective con-
example, Atlantic mackerel (Scomber scombrus) in
servation tool. There are no insurmountable scientific
southern England (Horwood et al. 1998). Further-
or technological hurdles to creating seamount pro-
more, (3) protection from fishing disturbance on
tected areas because they don’t move and their ben-
spawning grounds may increase spawning success.
thic and demersal species are mainly residents. If sea-
Nearshore spawning aggregations of northern cod
mount populations are largely self-recruiting, it could
(Gadus morhua) in Canada were trawled through 600
be easier to protect places to maintain and recover
to 1,880 times per year before the fishery was closed,
their species composition, structure, and ecosystem
and some shoals might have been trawled hundreds of
functions than in coastal waters, where species have
times per week at peak fishing intensities (Kulka et al.,
more open populations. In contrast, protecting
1995). Trawls disrupted aggregations for up to an hour
epipelagic species and habitats not associated with
after passage, and could have caused considerable but
fixed benthic features is more challenging because
unknown stress to fish (Morgan et al. 1997). (4) Indi-
pelagic hot spots are often ephemeral or shift posi-
rectly, migratory species could benefit from habitat
tions on short time scales, and because most of their
protection from damage by fishing gears in reserves.
large inhabitants are highly migratory. However, be-
Finally, (5) many supposedly migratory species appear
cause many pelagic species use predictable habitats to
to have more sedentary individuals that remain
MCBi.qxd
2/2/2005
2:41 PM
Page 313
Place-Based Ecosystem Management in the Open Ocean
313
within breeding grounds year-round and could bene-
fowl Habitat Areas of Major Concern (http://www
fit from fixed-location reserves.
.birds.cornell.edu/pifcapemay/pashleywarhurst.htm),
Nonmarine realms offer marine conservation biol-
including prairie potholes and other wetlands where
ogists important precedents for protecting migration
millions of ducks feed and nest. The combination of
routes and feeding, breeding, or nursery grounds. For
strong nesting habitat protection in some places and
instance, whooping cranes (Grus americana), which
well-regulated exploitation outside reserves led to re-
can migrate >4,000 km, had a very close brush with
covery of most duck populations and high, yet sus-
extinction due both to overexploitation and destruc-
tainable, levels of hunting.
tion of their wintering habitat (Lewis 1986). Estab-
Migratory corridors for terrestrial megafauna such
lishment of Wood Buffalo National Park, Northwest
as pronghorns (Antilocapra americana) often occupy
Territories (Canada), protected the crane’s breeding
very restricted, predictable portions of the landscape
grounds, while Aransas National Wildlife Refuge,
(Berger 2004). Protecting these areas has disporpor-
Texas (USA), protected remnants of their wintering
tionately high conservation benefit.
habitat. Whooping cranes spend most of their time in
Protection of spawning habitat and migratory corri-
reserves and are largely safe from hunting outside re-
dors used by heavily exploited migratory species is
serves thanks to protection under Canadian and US
hardly unfamiliar to marine scientists. A central reason
laws. The combination of conserving key habitats
for the survival of extant populations of Pacific salmon
and strong efforts that reduced mortality outside re-
(Hilborn, Chapter 15), some of which migrate thou-
serves has been essential to their recovery.
sands of kilometers, has been protection of streams
Highly migratory species do not need to spend a
where these fishes spawn and undergo early develop-
major fraction of their time in protected places to ful-
ment. Similarly, a number of nations protect localized
fill key conservation goals. Red knots (Calidris canutus)
herring spawning areas in temperate coastal regions to
and other shorebirds migrate as far as 10,000 km from
allow successful reproduction and egg survival. In trop-
breeding grounds on the Canadian tundra to Ar-
ical waters, snappers (Lutjanidae) and groupers (Ser-
gentina. Along the way they depend on a small num-
ranidae) often migrate tens or hundreds of kilometers
ber of Atlantic estuarine mudflats having the abun-
to spawning aggregation sites. Intensive fishing has ex-
dant food needed to replenish fat deposits. Because
tirpated many such aggregations throughout the trop-
loss of these areas would devastate shorebird popula-
ics (Sala et al. 2001), with consequential population-
tions, Manomet Center for Conservation Sciences or-
wide depletion for the species involved. Recognizing
ganized the Western Hemisphere Shorebird Reserve
that protection of spawning aggregations is essential for
Network of stepping-stone protected areas along
fishery sustainability, the US Virgin Islands seasonally
migration routes (http://www.manomet.org/WHSRN/
closes an aggregation site for red hind groupers (Epi-
history.htm). Although shorebirds spend only a brief
nephelus guttatus) to fishing. Although it only covers 1.5
time within the stopover and staging areas thereby
percent of the islands’ shelf fishing grounds, the closure
protected, this network is crucial to their conservation.
has led to populationwide increases in size of these
Heavily exploited highly migratory species can also
groupers (Beets and Friedlander 1999). Fisheries for
benefit from protection of key habitats. Most North
northern bluefin tuna in the Mediterranean have tra-
American ducks (Anatinae) are highly migratory. In
ditionally targeted them at bottlenecks where they mi-
the 1980s and ’90s, Canada, the United States, and
grate close to coasts and through narrow straits (Cush-
Mexico joined in the North American Waterfowl Man-
ing 1988). Protection of bottlenecks for this and other
agement Plan, a continent-scale effort to restore duck
pelagic species could offer them significant protection.
populations to their 1970s levels (Williams et al.
These examples clearly show that place-based con-
1999). The core of this effort is protecting 34 Water-
servation can be essential for conserving species that
MCBi.qxd
2/2/2005
314
2:41 PM
Page 314
Marine Conservation Biology
range far beyond protected area boundaries, including
umn could affect seamount demersal and benthic
species that use reserves for relatively brief periods and
communities. First, the longlines used at these sites
that are subjected to heavy human predation outside
average nearly 50 km in length, so lost gear armed
reserves. There is nothing about the open ocean that
with thousands of hooks could harm seafloor biota
would negate the efficacy of place-based conservation
through “ghost fishing.” Second and more impor-
so long as people don’t defeat its purpose by inter-
tant, the presence of fishing boats would greatly com-
cepting migrants as soon as they leave the protected
plicate some potentially potent monitoring and en-
area, a phenomenon called “fishing the line.”
forcement methods; radar satellite observations of
vessels might not be able to distinguish between per-
Benthic–Pelagic Coupling
mitted and illegal gear types (personal communica-
Place-based conservation aims to protect function-
tion with John Amos, SkyTruth, Shepardstown, West
ally intact ecosystems. The previous section addressed
Virginia, USA). Finally, as Australian government sci-
horizontal ecological linkages. Although vertical strat-
entists acknowledged, there are trophic linkages be-
ification is more important in open ocean ecosystems
tween seamounts’ benthic and pelagic communities
than on land, at least four key processes link various
via consumption of vertically migrating animals. Be-
depth strata: (1) primary production from the eu-
cause some large pelagic animals (e.g., swordfish,
photic zone rains organic particles to the seafloor; (2)
tunas, elephant seals, beaked whales) forage in the
diel vertical migrators transfer energy and nutrients
mesopelagic zone, fishing in overlying waters could
from the epipelagic zone to the depths; (3) diel verti-
affect seamount benthos. To our knowledge, there
cal migrators are eaten by epipelagic, mesopelagic,
has been no research addressing the role of large pred-
and seamount species; and (4) larvae of many benthic
ators in coupling pelagic and benthic seamount com-
species live, feed, and face risk of predation in the
munities, but there is one precedent from shallower
epipelagic zone until they settle. Despite these obvious
waters suggesting that this third consideration merits
vertical linkages, some supporters of protection of
study.
benthic communities on seamounts question whether
In experiments on Grand Bahama Bank (Bahamas),
there is any reason to protect pelagic species in their
Hixon and Carr (1997) found that there is strong den-
overlying waters. For example, Australia established
sity-dependent mortality in postsettlement resident
the Tasmanian Seamount Marine Reserve (http://www
planktivorous blue chromis (Chromis cyanea) only
.deh.gov.au/coasts/mpa/seamounts/index.html) in
when both resident piscivores (mostly groupers) and
1999 to protect about 70 untrawled seamounts that
visiting piscivores (mostly jacks, Carangidae) were
peak from 1,940 to 660 m below the surface. Bottom
present. Like some seamount fishes (e.g., rockfishes),
trawling, the fishing method that causes the most col-
blue chromis aggregations rise from the shelter of the
lateral damage (Chuenpagdee et al. 2003), is banned
spatially complex shallow coral fore reef matrix to
in this MPA, a sound management decision given the
feed in the water column. In the absence of transient
profound difference between untrawled and trawled
predators, predation on blue chromis by resident pred-
seamount ecosystems in this area (Koslow et al. 2001).
ators is relatively low. But when transient predators
But this MPA is not a true marine reserve: longlining
approach, blue chromis dive for cover among the
for southern bluefin tuna (Thynnus maccoyii) and
corals, making them more vulnerable to resident pred-
other large pelagic fishes, purse seining, squid jig-
ators. It would be surprising if the synergism between
ging, and noncommercial fishing are allowed in
these two guilds of coral reef predators did not operate
the water column as deep as 500 m over protected
on some seamounts (except, quite probably, with con-
seamounts.
sequences that play out much more slowly). The as-
There are several ways that fishing in the water col-
sumption that fishing in waters overlying seamounts is
MCBi.qxd
2/2/2005
2:41 PM
Page 315
Place-Based Ecosystem Management in the Open Ocean
O
O
FP
a.
315
FP
b.
F I G U R E 1 8 . 3 . Week-to-week changes in phytoplankton (chlorophyll a) concentration in oceanic ecosystems near
Hawaii (USA) based on MODIS Aqua ocean color images. Main Hawaiian islands are in black; light patches are areas
of higher chlorophyll a concentrations. 18.3a represents 25 January to 1 February 2004; 18.3b represents 2 to 9 February 2004 (NOAA Coast Watch Central Pacific Node Remote Sensing Archives, available at http://coastwatch.nmfs
.hawaii.edu/ocean_color/aqua_color_archive.html).
not a consequential management issue, and that fully
displacement of such key ecosystem features. Not
protected reserves are therefore unnecessary for sea-
only do place-based ecosystem managers have to over-
mounts, could benefit from critical examination.
come widespread lack of understanding that the open
ocean has places; they also have to deal with key eco-
Ecosystems That Move
system features that move and change shape.
On land, protecting places, however difficult, is not as
our terrestrially shaped minds. People fully under-
Monitoring, Compliance, and
Enforcement Issues
stand that there are places on land and are accustomed
Once, oceanic species and ecosystems were safe from
to dealing with maps, fences, and strong penalties for
anthropogenic harm because people were loath to
violating boundaries (EAN once saw a sign at a forest
venture too far offshore and fishing technologies
edge in North Carolina, saying “No trespassing. Sur-
could not find or reach animals that were too deep or
vivors will be prosecuted.”). But people have much
too fast. But the increasing adoption of new tech-
less sense of place in the ocean. A precondition for
nologies, especially in the last two decades, has dra-
place-based marine conservation is a broader under-
matically increased fishing power, eliminating de
standing that the sea actually has places with distinc-
facto refuges in the oceanic realm by turning the sea
tive attributes.
transparent and allowing fishermen to expand fishing
difficult as doing so in the sea. One reason comes from
On land, in the rocky intertidal, in shallow-water
grounds into the world’s remotest places, where they
coral reefs, and in oceanic ecosystems such as mido-
can fish on previously inaccessible habitats, including
cean ridges and seamounts, ecosystems don’t move at
rough, steep bottoms down to 2 km. Technology has
speeds humans can perceive. But some pelagic eco-
shifted the balance overwhelmingly against fishes.
systems can move kilometers or even tens of kilome-
But technology can also help to protect and recover
ters per day: particular isotherms, oxyclines, eddies,
oceanic ecosystems.
fronts, upwelling at divergence zones, and floating ob-
A number of existing technologies could be used
jects aggregated at convergence zones (Figure 18.3a
singly or together to ensure the effectiveness of
and 18.3b). Any effective pelagic reserve, MPA, or
oceanic fishery closures, MPAs, or fully protected re-
fishery closure will have to account for the appear-
serves. Even in the epipelagic zone, where fishes are
ance, disappearance, expansion, contraction, or lateral
not always easy to find and their habitats move, man-
MCBi.qxd
2/2/2005
316
2:41 PM
Page 316
Marine Conservation Biology
agers can determine important areas deserving pro-
high seas, a necessary complement to MPAs. Despite
tection by using new electronic tags to trace species’
these obstacles, there is growing hope that the inter-
movements and behavior. They can emulate fisher-
national community may be willing to take major
men by downloading daily images of oceanographic
steps within the next few years to enable this vision to
conditions known to attract particular species com-
become a reality (Gjerde 2003).
munities. Indeed, they can use data from researchers
The United Nations Convention on the Law of the
that correlate tagging data with oceanographic data so
Seas (UNCLOS, 1982) provides the legal basis for high
that specific oceanographic phenomena become a
seas MPAs, including oceanic reserves (Warner 2001).
proxy for the oceanic habitats used by fishes. Using
UNCLOS establishes unqualified obligations to protect
these data, managers could then set dynamic reserve
and preserve the marine environment and to cooper-
boundaries and broadcast the coordinates to fisher-
ate in conserving living resources. This includes the
men frequently, even daily.
specific obligation to take necessary measures “to pro-
Compliance of fishermen in nearshore areas is es-
tect and preserve rare or fragile ecosystems as well as
sential to the success of protected areas (Walmsley and
the habitat of depleted, threatened or endangered
White 2003), and this is even more true in the open
species and other forms of marine life” (Art. 194(5)).
ocean. Fortunately, oceanic fishermen, who tend to be
As there is general agreement that UNCLOS is so
ahead of enforcement personnel technologically, can
widely accepted among States that it also represents
readily determine their precise locations vis-à-vis
customary international law, its environmental obli-
those of closed areas by using global positioning sys-
gations apply to all States, not just those that have ex-
tems, which are commonplace in oceangoing vessels.
pressly agreed to adhere to the treaty (Scovazzi 2004).
Enforcement officials can use vessel monitoring sys-
The rights of States to exercise high seas “free-
tems (VMSs) and various kinds of tamper-proof event
doms” such as fishing, navigation, the laying of sub-
data recorders (EDRs), including video, to determine
marine cables and pipelines, marine scientific re-
the locations of fishing vessels relative to closed area
search, and construction of artificial islands and other
boundaries. VMSs and EDRs can also help to distin-
installations are subject to the conditions laid down in
guish innocent passage from illegal fishing, and offi-
UNCLOS, including the responsibility to protect and
cials can seek independent verification with radar
preserve the marine environment and conserve ma-
satellite images, which can “see” at night and through
rine living resources, as well as other rules of interna-
cloud cover, to detect vessels that enter reserves. As-
tional law (Art. 87) (Kimball 2004). Nevertheless, the
suring compliance with moving boundaries would
jurisdictional framework established in UNCLOS and
pose novel conceptual challenges, no doubt, but is
its lack of a well-developed system for enforcement
within easy technological reach.
with respect to high seas environmental and conservation obligations means that there are few mecha-
High Seas Jurisdiction
nisms to assist States who want to protect high seas
Establishing networks of oceanic reserves, other MPAs,
biodiversity. The present system leaves many oppor-
and fishery closures on the high seas will be a major
tunities for States to avoid taking responsibility for
challenge under current international law. Though
their own action or that of their citizens or “flag”
the legal basis exists for sovereign States to agree to
ships registered in their country.
protect discrete areas of the high seas environment,
In waters subject to national jurisdiction (out to
there are three major obstacles: (1) the need to secure
200 nautical miles from shore) and on their legally de-
the agreement of affected nations, (2) the difficulty of
fined outer continental shelf when it extends beyond
enforcing regulations, and (3) the lack of an adequate
200 nm (set by a complex formula under UNCLOS
framework for ecosystem-based management of the
Article 76), coastal States may regulate the resource-
MCBi.qxd
2/2/2005
2:41 PM
Page 317
Place-Based Ecosystem Management in the Open Ocean
317
related activities of their own citizens and foreign na-
regional adoption of “Specially Protected Areas of
tionals. They may establish and enforce protected
Mediterranean Importance” on the high seas. The
areas for these activities. Within 200 nm, coastal States
Mediterranean States have already agreed to recognize
also have broad powers of environmental protection,
and protect the Pelagos Sanctuary for Mediterranean
balanced with rights of other States to navigation and
Marine Mammals in the Ligurian Sea, 50 percent of
overflight, and the laying of submarine cables and
which is in international waters. The OSPAR Conven-
pipelines (UNCLOS Art. 58). But beyond national ju-
tion for Protection of the Northeast Atlantic also fore-
risdiction the situation is very different. On the high
sees establishment of protective measures, including
seas beyond 200 nm (including above their outer con-
MPAs, throughout the maritime area covered by the
tinental shelf), States may only regulate the behavior
agreement, a large part of which is high seas.
of their nationals and ships flagged under their laws.
Thus regional agreements between likeminded
Moreover, “flag States” are the only States with au-
States can provide a high degree of recognition and
thority to enforce regulations against their ships while
protection to discrete high seas areas, based on the
on the high seas, unless otherwise agreed by the States
willingness of the States to police their own citizens
concerned. Thus, despite strong environmental and
and ships (Young 2003). However, these regional
conservation obligations under international law to
agreements must still secure widespread endorsement
refrain from damaging activities, each State must
and compliance. In Antarctica, the consent of fishing
agree to regulate its nationals and ships regarding ac-
States that are members of the Commission for the
tivities that may affect a high seas MPA or reserve. It
Conservation of Antarctic Marine Living Resources
is therefore important to involve at least all the States
(CCAMLR) must first be obtained before a marine
whose activities may affect the site. These States may
ASPA is established. In the Mediterranean, the con-
act individually or decide to enter into an agreement:
cerned States have realized they have little ability to
for example, States that are members of a regional
control illegal actions of vessels from outside the re-
fishery management organization (RFMO) may agree
gion affecting the Pelagos Sanctuary and its protected
to close a particular area to fishing. States seeking to
cetaceans. In the northeast Atlantic, the States have
establish an oceanic reserve therefore must convince
ceded responsibility to regulate fisheries to the Euro-
other nations that the benefits of protecting these eco-
pean Union and the Northeast Atlantic Fisheries Com-
systems outweigh the short-term costs and inconven-
mission, and thus have agreed not to address fisheries
ience, in addition to legal arguments.
impacts under the OSPAR Convention. To have effec-
The good news is that States have already reached
tive networks of areas protected from harmful activi-
agreement to establish highly protected areas in inter-
ties, a high level of international cooperation and
national waters in several important regions. The Pro-
consistent application of ecosystem-based and pre-
tocol on Environmental Protection to the Antarctic
cautionary approaches will be essential. More detailed
Treaty (1991) envisages the development of a full range
agreements and appropriate mechanisms may be
of “Antarctic Specially Protected Areas,” including
needed.
marine areas, to protect “outstanding environmental,
Developments in international fisheries law may be
scientific, historic, aesthetic, or wilderness values, or
of prime importance in the future evolution of high
ongoing or planned scientific research.” Entry is pro-
seas closures, MPAs, and reserves. The 1995 UN Agree-
hibited without a permit. There are now six fully ma-
ment on Straddling Fish Stocks and Highly Migratory
rine “Antarctic Specially Protected Areas” (ASPAs) in the
Fish Stocks (UN Fish Stock Agreement [FSA]) was de-
Southern Ocean. The Protocol on Specially Protected
veloped to implement the provisions of UNCLOS with
Areas and Biodiversity to the Barcelona Convention for
respect to fish stocks crossing national and interna-
the Mediterranean (1995) has provisions to enable the
tional waters. Articles 5 and 6 contain significant en-
MCBi.qxd
2/2/2005
318
2:41 PM
Page 318
Marine Conservation Biology
vironmental obligations, calling for States party to the
agreed conservation and management measure that
agreement to (1) minimize the impact of fishing on
must be complied with to gain access to the fish stock;
nontarget, associated, and dependent species, and eco-
support from key fishing nations; and strong enforce-
systems; (2) protect habitats of special concern; (3)
ment measures. However three weaknesses remain
apply the precautionary approach; and (4) protect bio-
obstacles to using the UN FSA as a vehicle for net-
diversity. Thus it clearly envisages areas closed to fish-
works of high seas reserves and MPAs:
ing activities, not just for the management of fish
stocks but also for the protection of biological diversity.
The UN FSA is also a significant precedent because
1. Only 51 States are parties to the UN FSA, though
this now includes all 15 members of the Euro-
of its provisions clarifying the duty of States party to
pean Union, the United States, Russia, Norway,
the agreement to cooperate to conserve high seas liv-
India, Brazil, Ukraine, and several South Pacific
ing resources and enabling nonflag States to enforce
nations.
conservation measures. The UN FSA requires States to
2. It only applies to fish stocks that straddle na-
collaborate through regional fisheries management
tional and international waters and to highly
organizations or less formal arrangements. States
migratory fish stocks such as tunas and swordfish,
wishing to fish on a stock already subject to agreed
and not to discrete high seas fish stocks, such as
conservation and management measures must either
orange roughy dwelling on midocean seamounts.
join the relevant RFMO or arrangement, or must abide
3. It is only slowly being incorporated by regional
by the applicable conservation measures. It further re-
fisheries management organizations, the
quires flag States to exercise serious oversight and
implementing arm for this agreement.
control over its vessels authorized to fish on the high
seas (de Fontaubert and Luchtman 2003).
Moreover, agreements pursuant to the UN FSA would
To encourage compliance, the UN FSA recognizes
not be able to control other potentially harmful activ-
the right of nonflag States to take enforcement action
ities. This would require petitioning other interna-
against foreign vessels on the high seas. This is still
tional agencies, such as the International Maritime Or-
couched in very moderate language within complex
ganization for regulations to prevent shipping impacts,
formulas to prevent abuse, but it marks a significant
and the International Seabed Authority for protection
recognition by States of the need to respond to lax
from mining activities. Certain key mechanisms com-
compliance and poor enforcement by flag States in
monly used in national waters to provide a framework
the past. It also recognizes the right of a port State
for sustainable management outside of protected areas,
(when a fishing vessel voluntarily enters its ports) to
such as environmental impact assessments and spatial
enforce agreed conservation measures. Recent ad-
planning/zoning would still be lacking.
vances in satellite remote sensing and global posi-
Nevertheless, there is reason to hope. Recent events
tioning technology, coupled with a heightened naval
have shown that the international community has fi-
interest and involvement, means that earlier problems
nally recognized the need to conserve and sustainably
of monitoring vessel activities may soon be a thing of
use high seas biodiversity and is committed to taking
the past.
action (Kimball 2004). At the World Summit on Sus-
High seas reserves, MPAs, and fishery closures es-
tainable Development in Johannesburg in 2002, world
tablished by a regional fisheries management organi-
leaders agreed to “maintain the productivity and bio-
zation that fully reflect the principles and provisions
diversity of important and vulnerable marine areas,
of the UN FSA would thus contain several of the key
including in areas beyond national jurisdiction.” They
ingredients necessary for successful management and
further called for adoption of the ecosystem-based ap-
protection: recognition of the protected area as an
proach in the marine environment, the elimination of
MCBi.qxd
2/2/2005
2:41 PM
Page 319
Place-Based Ecosystem Management in the Open Ocean
319
destructive fishing practices, and the establishment of
two kinds of protected places, ones that conserve bio-
MPAs, including representative networks by 2012 and
diversity in features that don’t move and others that
time/area closures for the protection of nursery
focus on features that move on timescales as short as
grounds and periods. Such protected areas, they un-
days.
derscored, must be consistent with international law
and based on scientific information.
Protecting Ecosystems with Fixed Boundaries
The United Nations General Assembly (UNGA) has
Many seamounts, submerged mountain ranges, and
endorsed these commitments and further called for
submarine canyons have benthic communities dom-
urgent consideration of ways to improve manage-
inated by deep-sea corals and sponges, which scien-
ment of risks to marine biodiversity on seamounts,
tists increasingly consider high conservation priorities
cold-water coral reefs, and certain other underwater
(MCBI 2004). Tunas, billfishes, marine mammals,
features. For areas beyond national jurisdiction, the
seabirds, and sea turtles often congregate in waters
UNGA has invited relevant international and regional
above these features where they can forage on
organizations to consider urgently how to better ad-
epipelagic or vertically migrating fishes, crustaceans,
dress risks, on a scientific and precautionary basis,
or squids. Protecting these places can safeguard the
how to use existing treaties and other relevant instru-
most vulnerable benthic communities and allow large
ments to do so, to identify marine ecosystem types
pelagic fishes to spawn or feed where their aggrega-
that warrant priority attention, and to explore a range
tions would otherwise make them most vulnerable to
of potential approaches and tools for their protection
fishing. Static features are the easiest oceanic places to
and management (UNGA Res.A/58/240). The States
protect because their locations can readily be identi-
party to the Convention on Biological Diversity have
fied by fishermen and enforcement officials alike. To
also expressed their concern and called for urgent ac-
be effective, however, establishing the boundaries for
tion (Dec. VII/5, paras. 61–62). Thus we can expect to
these areas may well require ecological understanding
see significant progress on these issues over the next
that goes beyond drawing lines around the features.
several years.
As Hooker et al. (2002) note, surrounding areas may
It is clear that support from the scientific commu-
provide crucial food subsidies to species that forage
nity will be essential to developing the scientific basis
above fixed features, so incorporating protected buffer
for place-based conservation in the open ocean. Just as
areas around features to be protected will provide
important, the support of the scientific community is
needed insurance for fixed protected ecosystems in
essential to educate policy makers on the importance
the open ocean.
and significance of high seas biological diversity and
ally agreed action to provide for ecosystem-based and
Protecting Ecosystems with
Dynamic Boundaries
precautionary management.
Although terrestrial ecosystems can undergo dramatic
ecosystem processes, and of the need for internation-
seasonal or episodic shifts in vegetation phenology
Options for Conserving Ecosystems in
the Open Ocean
and animal populations, protected places on land
have fixed boundaries, reflecting humans’ relatively
static concept of place in the terrestrial realm. So do
Networks of fishery closures, MPAs, and fully pro-
existing MPAs. But, as explained earlier, many key
tected reserves in the open ocean, whether within na-
habitats for large pelagic species shift or appear and
tions’ EEZs or on the high seas, will have to reflect
disappear with changing oceanographic conditions.
new understanding, new opportunities, and new
There are two ways for decision makers to conserve
kinds of international cooperation. We can envision
life in dynamic pelagic hot spots. One is to protect
MCBi.qxd
2/2/2005
320
2:41 PM
Page 320
Marine Conservation Biology
areas large enough to encompass shifting or ephem-
kinds of commercial, subsistence, or recreational fish-
eral habitats wherever they are. MPAs of enormous
ing), and others in permanent MPAs and reserves. The
size are not unprecedented: The Indian Ocean Whale
central question in achieving an acceptable balance is,
Sanctuary established by the International Whaling
How can managed areas protect enough of the open
Commission in 1979 prohibits whaling throughout
ocean to maintain and recover biodiversity to accept-
the Indian Ocean north of 55°S. But this is a single-
able levels without making so much off limits that fish-
purpose MPA; fishing for other marine life is allowed
eries cannot be profitable and productive in the short
there. Fully protected reserves of that scale seem eco-
term? A promising ecosystem-based management tool
nomically and politically unfeasible for the foresee-
to do this is creating networks of protected areas.
able future. An alternative—one that requires more
Networks are collections of protected places having
creative thinking—is to protect shifting habitats as
the emergent property of conserving more effectively
they move, reflecting decadal, interannual, seasonal,
as a whole than they would if their protected places
lunar, or even daily shifts in pelagic habitats. Of
were not connected. Consider an oceanic species
course, designing such fishery closures, MPAs, or re-
whose single population feeds only in one place,
serves with flexible boundaries requires comprehen-
breeds only in another, and migrates between them.
sive observations of changing ocean conditions, sci-
Without knowing where and when it feeds, breeds,
entific understanding about habitat needs, and the
and moves between these areas, one way to conserve
wealth of understanding that fishermen have gained
it is permanently protecting enough to be certain
through experience and observation. New tagging
that the area includes all of the above. An alternative
technologies and satellite observation of ocean con-
that can put far less of the ocean off limits, however,
ditions make this possible for the first time.
is to protect only its actual feeding area, breeding area,
Bluefin tuna schools, for example, aggregate along
and the migration route that connects them. Indeed,
temperature fronts in the Gulf of Maine that separate
some of these protections can be temporary. Of
warm and cooler water masses. Fishermen use satellite
course, this approach requires meaningful quantita-
temperature maps to help find these schools. The
tive knowledge of where these places are and when
same temperature data they use could also be used by
they are occupied by the species communities of con-
managers to set protected area boundaries—even on
cern. Moreover, the foregoing example constitutes
a day by day basis—and the coordinates of the pro-
the simplest of networks. As the number of places and
tected areas broadcast to the fishing fleet. There is no
interconnections increase, as occurs in species that
legal or technological reason why protected areas with
have metapopulation structure (Lipcius et al., Chapter
dynamic boundaries cannot be used to protect pelagic
19), networks must become more complex to reflect
species.
the biological realities in a dynamic ocean. Networks,
then, are ways to provide near-maximum conserva-
Networking Reserves and Other PlaceBased Ecosystem Management Areas
tion benefits while reducing regulation consistent
with achieving conservation objectives.
Network designs for seamounts and epipelagic spe-
Just as fishermen want the freedom to fish every-
cies are likely to be different. If seamount species were
where, people who want to conserve biodiversity
all self-recruiting, networks would be unnecessary be-
would like to protect everything. If society expands its
cause individual reserves that protect each seamount
interest in the oceans beyond producing meat to
would be sufficient; there would be no need for con-
maintaining marine biodiversity, any workable solu-
nectivity. But that level of genetic and ecological isola-
tion will be somewhere in between, with some places
tion is unlikely even for the most isolated seamounts.
always or sometimes open for fishing (or to particular
As recruitment, food organisms, and predators from
MCBi.qxd
2/2/2005
2:41 PM
Page 321
Place-Based Ecosystem Management in the Open Ocean
321
other locations become more important, networking is
the open ocean, but are likely to be key components
all the more necessary to maintain the connectivity and
of any comprehensive strategy for conserving oceanic
long-term viability of these populations.
species and ecosystems. As on land, what happens
Of course, seamounts host both benthic and
outside these sanctuaries is crucial (Allison et al. 1998;
pelagic species. It is conceivable that their pelagic
Baum et al. 2003; Crowder et al. 2000; Lipcius et al.,
species could be resident, which would imply that
Chapter 19). Simply displacing fishing effort —
connectivity needs for seamount benthic species
thereby increasing fishing in unprotected areas—is
would be suitable to conserve their pelagics as well.
likely to have unwelcome results. Reducing fishing ef-
But if their pelagic species are highly migratory, as
fort is essential.
seems likely, then seamount ecosystem-based man-
Effective place-based management of oceanic spe-
agement will need to address the distinctive needs of
cies will require population and ecosystem research
pelagic visitors. In other words, oceanic ecosystem-
and monitoring to help managers set ecologically sus-
based managers will have to become as sophisticated
tainable catch limits. It will require precautionary de-
as those who manage shorebirds, ducks, and salmon.
cision making that makes protecting and recovering
Similarly, for dynamic and ephemeral pelagic fea-
biodiversity the highest priority. Effective means of as-
tures (e.g., fronts, upwellings) whose temporary ag-
suring compliance, including using off-the-shelf and
gregations of migratory wildlife need protection,
new technologies to monitor fishing operations and
maintaining connectivity is a central conservation
effective enforcement, are also essential. These, in
requirement, so networking becomes the primary con-
turn, will necessitate both creative new thinking
servation tool. Reserves are clearly the most foolproof
about managing marine ecosystems and adequate
tools for assembling effective networks. MPAs that
funding. But given the declining populations of sea-
provide less than full protection are likely to have
mount species and pelagic megafauna, it is clear that
greater enforcement costs, and temporary fishery clo-
existing management measures, by themselves, are
sures are more costly still because they require more
not adequate. New ecosystem-based approaches, in-
up-to-date information on habitat use. Where moni-
cluding networks that protect key places, are needed
toring and management systems are highly effective,
to change declining population trajectories of oceanic
networks can be assembled by establishing intercon-
animals.
nected reserves and less protected areas, or ocean zoning (see Norse, Chapter 25).
Conclusions
Acknowledgments
The idea for this chapter arose at a scientific workshop
titled “Ecology and Conservation Biology of Large
Ecosystem-based management that protects oceanic
Pelagic Fishes” held by Marine Conservation Biology
fixed habitats, such as seamounts, and shifting fea-
Institute and The Billfish Foundation in Islamorada,
tures, such as fronts, is a new idea, but one with
Florida (USA), in 1997, when Hazel Oxenford and Rus-
many relevant precedents in terrestrial and nearshore
sell Nelson suggested an examination of ways that
marine realms. The legal basis for reserves in interna-
protected areas could be used for conserving billfishes
tional waters already exists, but new institutional
and large tunas. We thank them and all the workshop
arrangements are needed and the viability of place-
participants, as well as the Offield Family Foundation
based conservation remains, so far, untested. We be-
and the William H. and Mattie Wattis Harris Founda-
lieve that networks of interconnected fishery closures,
tion, who funded the workshop. We thank the Pew
partly protected MPAs, and fully protected marine re-
Charitable Trusts for funding research by LBC on the
serves will not be the only tools for conservation in
global impact of pelagic longline fisheries and the Pew
MCBi.qxd
2/2/2005
322
2:41 PM
Page 322
Marine Conservation Biology
Fellows in Marine Conservation for supporting KMG’s
and the decline of pelagic sharks in the Gulf of
work on high seas governance. We are grateful to the
Mexico. Ecology Letters 7: 135–145
J.M. Kaplan Fund and the Oak Foundation for funding
Baum, J.K., R.A. Myers, D.G. Kehler, B. Worm, S.J.
MCBI’s work on seamount conservation and to the
Harley, and P.A. Doherty (2003). Collapse and con-
J.M. Kaplan Fund for funding IUCN’s work on high
servation of shark populations in the Northwest At-
seas conservation. We thank Lance Morgan, Fan Tsao,
lantic. Science 299: 389–392
Sara Maxwell, and Katy Balatero of MCBI; Rebecca
Beets, J. and A. Friedlander (1999). Evaluation of a
Lewison, Sloan Freeman, and Caterina D’Argrosa of
conservation strategy: A spawning aggregation clo-
Duke University Marine Laboratory; John Amos of
sure for red hind, Epinephelus guttatus, in the US Vir-
SkyTruth; and Paul Dayton and Karin Forney, for pro-
gin Islands. Environmental Biology of Fishes 55: 91–
viding stimulating ideas and information, and Masa
98
Ushioda of Coolwater Photos for his oceanic whitetip
Berger, J. (2004). The last mile: How to sustain long-
shark image. Finally, we thank Lee Kimball for her in-
distance migration in mammals. Conservation Biol-
sightful comments on the legal section, and Claudia
ogy 18(2): 320–331
Mills and John Twiss for their ideas and constructive
criticism of early drafts.
Block, B.A., D.P. Costa, G.W. Boehlert, and R.E.
Kochevar (2002). Revealing pelagic habitat use: The
tagging of Pacific pelagics program. Oceanologica
Literature Cited
Acta 25: 255–266
Boehlert G.W. and A. Genin (1987). A review of the ef-
Agardy, T.S. (1997). Marine Protected Areas and Ocean
fects of seamounts on biological processes. Pp. 319–
Conservation. R.G. Landes Company, Austin, Texas
334 in B.H. Keating, P. Fryer, R. Batiza, and G.W.
(USA)
Boehlert. eds. Seamounts, Islands and Atolls. Geo-
Allison, G.W., J. Lubchenco, and M.H. Carr (1998).
Marine reserves are necessary but not sufficient for
marine conservation. Ecological Applications 8(1)
Supplement: S79–S92
Angel, M.V. (1993). Biodiversity of the pelagic ocean.
Conservation Biology 7(4): 760–772
physical Monograph 43. American Geophysical
Union, Washington, DC (USA)
Botsford, L.W., F. Micheli, and A. Hastings (2003).
Principles for the design of marine reserves. Ecological Applications 13(1) Supplement: S25–S31
Branch, T.A. (2001). A review of orange roughy Ho-
Apostolaki, P., E.J. Milner-Gulland, M.K. McAllister,
plostethus atlanticus fisheries, estimation methods,
and G.P. Kirkwood (2002). Modelling the effects of
biology and stock structure. South African Journal of
establishing a marine reserve for mobile fish species.
Canadian Journal of Fisheries and Aquatic Sciences
59(3): 405–415
Marine Science 23: 181–203
Bull, B., I. Doonan, D. Tracey, and A. Hart (2001). Diel
variation in spawning orange roughy (Hoplostethus
Auster, P. and R. Langton (1999). The effects of fishing
atlanticus, Trachichthyidae) abundance over a sea-
on fish habitat. Pp. 150–187 in L.R. Benaka, ed. Fish
mount feature on the northwest Chatham Rise.
Habitat: Essential Fish Habitat and Rehabilitation.
New Zealand Journal of Marine and Freshwater Re-
American Fisheries Society, Symposium 22. American Fisheries Society, Bethesda, Maryland (USA)
search 35 (3): 435–444
Cailliet, G.M., A.H. Andrews, E.J. Burton, D.L. Watters,
Auster, P.J., J. Lindholm, and P.C. Valentine (2003).
E.E. Kline, and L.A. Ferry-Graham (2001) Age de-
Variation in habitat use by juvenile Acadian redfish,
termination and validation studies of marine fishes:
Sebastes fasciatus. Environmental Biology of Fishes 68:
Do deep-dwellers live longer? Experimental Geron-
381–389
tology 36: 739–764
Baum, J.K. and R.A. Myers (2004). Shifting baselines
Carlton, J.T., J.B. Geller, M.L. Reaka-Kudla, and E.A.
MCBi.qxd
2/2/2005
2:41 PM
Page 323
Place-Based Ecosystem Management in the Open Ocean
323
Norse (1999). Historical extinctions in the sea. An-
posed rule. Federal Register 65 (August 1): 47214–
nual Review of Ecology and Systematics 30: 515–538
47238
Christensen, V., S. Guénette, J.H. Heymans, C.J. Wal-
Department of Commerce (2001). Atlantic highly mi-
ters, R. Watson, D. Zeller, and D. Pauly (2003).
gratory species: Pelagic longline fishery; sea turtle
Hundred-year decline of North Atlantic predatory
protection measures. Federal Register 66 ( July 13):
fishes. Fish and Fisheries 4: 1–24
36711–36714
Chuenpagdee, R., L.E Morgan, S.M Maxwell, E.A
Druffel, E.R.M., S. Griffin, A. Witter, E. Nelson, J.
Norse, and D. Pauly (2003). Shifting gears: Assessing
Southon, M. Kashgarian, and J. Vogel (1995). Ger-
collateral impacts of fishing methods in US waters.
ardia: Bristlecone pine of the deep sea? Geochimica
Frontiers in Ecology and the Environment 1(10): 517–
et Cosmochimica Acta 59: 5031–5036
524
Conover, D.O., J. Travis, and F.C. Coleman (2000). Essential fish habitat and marine reserves: An intro-
Dulvy, N.K., Y. Sadovy and J.D. Reynolds (2003). Extinction vulnerability in marine populations. Fish
and Fisheries 4(1): 25–64
duction to the Second Mote Symposium in Fisheries
Estes, J.A. and J.F. Palmisano (1974). Sea otters: Their
Ecology. Bulletin of Marine Science 66(3): 527–534
role in structuring nearshore communities. Science
Crooks, K.R. and M.E. Soulé (1999). Mesopredator release and avifaunal extinctions in a fragmented system. Nature 400: 563–566
Crowder, L.B., S.J. Lyman, W.F. Figueira, and J. Priddy
(2000). Source-sink population dynamics and the
problem of siting marine reserves. Bulletin of Marine
Science 66: 799–820
Cushing, D.H. 1988. The Provident Sea. Cambridge
University Press, Cambridge (UK)
185 (4156): 1058–1060
Estes, J A., M.T. Tinker, T.M. Williams, and D.F. Doak
(1998). Killer whale predation on sea otters linking
oceanic and nearshore ecosystems. Science 282
(5388): 473–476
Food and Agriculture Organization (FAO) (2002). The
State of World Fisheries and Agriculture, 2002. UN
Food and Agriculture Organization, Rome (Italy)
Food and Agriculture Organization (FAO) (2003). Fish-
Dayton, P.K. (1975). Experimental studies of algal
eries Management, 2: The Ecosystem Approach to Fish-
canopy interactions in a sea otter–dominated kelp
eries. UN Food and Agriculture Organization, Rome
community at Amchitka Island, Alaska. Fishery Bul-
(Italy). Available at www.fao.org/DOCREP/005/
letin 73: 230–237
Y4470E/Y4470E00.HTM
Dayton, P.K., S.F. Thrush, M.T. Agardy, and R.J. Hof-
Fickling, D. (2003). Antarctic chase ends with arrests:
man (1995). Environmental effects of marine fish-
Australian and South African officials board suspect
ing. Aquatic Conservation: Marine and Freshwater Eco-
fishing boat. Guardian, August 28, 2003. Available at
systems 5: 205–232
www.guardian.co.uk/fish/story/0,7369,1030608,00
de Fontaubert, C. and I. Luchtman (2003). Achieving
.html
Sustainable Fisheries: Implementing the New Inter-
Gell F.R. and C.M. Roberts (2003). Benefits beyond
national Legal Regime. IUCN, Gland (Switzerland)
boundaries: The fishery effects of marine reserves.
and Cambridge (UK). Available at www.iucn.org/
Trends in Ecology and Evolution 18: 448–455
themes/marine
Genin, A., C. Greene, L. Haury, P. Wiebe, G. Gal, S.
Department of Commerce (1999). Western Pacific
Kaartvedt, E. Meir, C. Fey, and J. Dawson (1994).
pelagic fisheries: Hawaii-based pelagic longline area
Zooplankton patch dynamics: Daily gap formation
closure. Federal Register 64 (December 27): 72290–
over abrupt topography. Deep Sea Research I 41:
72291
941–951
Department of Commerce (2000). Atlantic highly mi-
Gislason, H., M. Sinclair, K. Sainsbury, and R. O’Boyle
gratory species: Pelagic longline management: pro-
(2000). Symposium overview: Incorporating ecosys-
MCBi.qxd
2/2/2005
324
2:41 PM
Page 324
Marine Conservation Biology
tem objectives within fisheries management. ICES
Energy demand of foraging bottlenose whales (Hy-
Journal of Marine Science 57: 468–475
peroodon ampullatus) in a marine protected area. Bi-
Gjerde, K.M., ed. (2003). Ten-year strategy to promote
ological Conservation 104: 51–58
the development of a global representative system
Horwood, J.W., J.H. Nichols, and S. Milligan (1998).
of high seas marine protected area networks, as
Evaluation of closed areas for fish stock conserva-
agreed by Marine Theme Participants at the 5th
tion. Journal of Applied Ecology 35: 893–903
World Parks Congress Governance Session “Pro-
Houde, E., ed. (2001). Marine Protected Areas: Tools for
tecting Marine Biodiversity beyond National Juris-
Sustaining Ocean Ecosystems. National Academy
diction,” Durban, South Africa (8–17 September
2003). IUCN, WCPA WWF. Available at www.iucn
.org/themes/marine/pdf/10ystrat.pdf
Gjerde, K.M. and C. Breide (2003). Towards a Strategy
for High Seas Marine Protected Areas: Proceedings of the
Press, Washington, DC (USA)
Hunt, G. L., Jr., K.O. Coyle, S. Hoffman, M.B. Decker,
and E.N. Flint (1996). Foraging ecology of shorttailed shearwaters near the Pribilof Islands, Bering
Sea. Marine Ecology Progress Series 141: 1–11
IUCN, WCPA and WWF Experts Workshop on High
Hyrenbach, K.D., P. Fernández, and D.J. Anderson
Seas Marine Protected Areas, 15–17 January 2003,
(2002). Oceanographic habitats of two sympatric
Malaga, Spain. IUCN, Gland (Switzerland)
North Pacific albatrosses during the breeding sea-
Goñi, R., N.V.C. Polunin, and S. Planes (2000). The
son. Marine Ecology Progress Series 233: 283–301
Mediterranean: Marine protected areas and the re-
Hyrenbach, K.D., K.A. Forney, and P.K. Dayton (2000).
covery of a large marine ecosystem. Environmental
Marine protected areas and ocean basin manage-
Conservation 27: 95–97
ment. Aquatic Conservation: Marine and Freshwater
Gulf of Mexico Fishery Management Council (1999).
Ecosystems 10: 437-458
Marine Reserves Technical Document: A Scoping Docu-
Jackson, J.B.C., M.X. Kirby, W.H. Berger, K.A. Bjorndal,
ment for the Gulf of Mexico. Gulf of Mexico Fisheries
L.W. Botsford, B.J. Bourque, R.H. Bradbury, R.
Management Council, Tampa, Florida (USA)
Cooke, J. Erlandson, J.A. Estes, T.P. Hughes, S. Kid-
Haury, L., C. Fey, G. Gal, A. Hobday, and A. Genin
well, C.B. Lange, H.S. Lenihan, J.M. Pandolfi, C.H.
(1995). Copepod carcasses in the ocean, I: Over
Peterson, R.S. Steneck, M.J. Tegner, and R.R. Warner
seamounts. Marine Ecology Progress Series 123: 57–63
(2001). Historical overfishing and the recent col-
Haury L., C. Fey, C. Newland, and A. Genin (2000).
lapse of coastal ecosystems. Science 293: 629–638
Zooplankton distribution around four eastern
Juda, L. (2003). Changing national approaches to
North Pacific seamounts. Progress in Oceanography
ocean governance: The United States, Canada, and
45 (1): 69–105
Australia. Ocean Development and International Law
Henke, S.E. and F.C. Bryant (1999). Effects of coyote
removal on the faunal community in western Texas.
Journal of Wildlife Management 63: 1066–1081
Hixon, M.A. and M.H. Carr (1997). Synergistic predation, density dependence, and population regulation in marine fish. Science 277: 946–949
34(2): 161–187
Kimball, L. (2004). International Conservation Initiatives. Paper prepared for AAAS Annual Meeting,
Seattle, Washington, 12–16 February 2004
Klimley A.P. (1995). Hammerhead City. Natural History
104 (10): 32–38
Hooker, S.K. and L.R. Gerber (2004). Marine reserves as
Koslow, J.A., G.W. Boehlert, J.D.M. Gordon, R.L.
a tool for ecosystem-based management: The po-
Haedrich, P. Lorance, and N. Parin (2000). Conti-
tential importance of megafauna. BioScience 54 (1):
nental slope and deep-sea fisheries: Implications for
27–39
a fragile ecosystem. ICES Journal of Marine Science 57:
Hooker, S.K., H. Whitehead, and S. Gowans (2002).
Ecosystem consideration in conservation planning:
548–557
Koslow, J.A., K. Gowlett-Holmes, J.K. Lowry, T. O’Hara,
MCBi.qxd
2/2/2005
2:41 PM
Page 325
Place-Based Ecosystem Management in the Open Ocean
325
G.C.B. Poore, and A. Williams (2001). Seamount
ford, M.H. Carr, G.E. Davis, P.K. Dayton, D. Got-
benthic macrofauna off southern Tasmania: Com-
shall, D.R. Gunderson, M.A. Hixon, J. Lubchenco,
munity structure and impacts of trawling. Marine
M. Mangel, A. MacCall, D.A. McArdle, J.C. Ogden,
Ecology Progress Series 213: 111–125
J. Roughgarden, R.M. Starr, M.J. Tegner, and M.M.
Kulka, D.W., J.S. Wroblewski, and S. Narayanan (1995).
Yoklavich (1999). No-take reserve networks: Pro-
Recent changes in the winter distribution and move-
tection for fishery populations and marine eco-
ment of the northern Atlantic cod (Gadus morhua
Linnaeus 1758) on the Newfoundland–Labrador
shelf. ICES Journal of Marine Science 52: 889–902
Lewis, J.C. (1986). The whooping crane. Pp. 659–676
in R.L. Di Silvestro, ed. Audubon Wildlife Report
systems. Fisheries 24:11–25
Musick, J.A., ed. (1999). Life in the Slow Lane: Ecology
and Conservation of Long-Lived Marine Animals.
American
Fisheries
Society
Symposium,
23.
Bethesda, Maryland (USA)
1986. National Audubon Society, New York (USA)
Myers, R.A. and B. Worm (2003). Rapid worldwide de-
Littler, M.M., D.S. Littler, S.M. Blair, and J.N. Norris
pletion of predatory fish communities. Nature 423:
(1985). Deepest known plant life discovered on an
uncharted seamount. Science 227: 57–59
280–283
Norse, E.A. (2003). Why marine protected areas? Pp.
Lubchenco, J., S.R. Palumbi, S.D. Gaines, and S. An-
1–9 in J.P. Beumer, A. Grant, and D.C. Smith, eds.,
delman (2003). Plugging a hole in the ocean: The
Aquatic Protected Areas: What Works Best and How Do
emerging science of marine reserves. Ecological Ap-
We Know? Proceedings of the World Congress on
plications 13(1) Supplement: S3–S7
Aquatic Protected Areas, 14–17 August, 2002, Cairns,
Lueck, R.G. and T.D. Mudge (1997). Topographically
induced mixing around a shallow seamount. Science
276 (5320): 1831–1833
Marine Conservation Biology Institute (MCBI) (2004).
Scientists’ Statement on Protecting the World’s Deep-Sea
Australia. University of Queensland Printery, St.
Lucia, Queensland (Australia)
Paine, R.T. (1966). Food web complexity and species
diversity. American Naturalist 100: 65–75
Paine, R.T. (1969). A note on trophic complexity and
Coral and Sponge Ecosystems. Statement signed by
species diversity. American Naturalist 103: 91–93
1,136 scientists, released at the Annual Meeting of
Parrish, R. (1999). Marine reserves for fisheries man-
the American Association for the Advancement of
agement: Why not. California Cooperative Oceanic
Science, Seattle, Washington (USA). Available at
Fisheries Investigations Reports 40: 77–86
www.mcbi.org/DSC_statement/sign.htm
Merrett, N.R. and R.L. Haedrich (1997). Deep-Sea Demersal Fish and Fisheries. Chapman & Hall, London
(UK)
Mills, C.E. and J.T. Carlton (1998). Rationale for a system of international reserves for the open ocean.
Conservation Biology 12(1): 244–247
Mooney, H., ed. (1999). Sustaining Marine Fisheries. National Academy Press, Washington, DC (USA)
Pauly, D., V. Christensen, J. Dalsgaard, R. Froese, and
F. Torres (1998). Fishing down marine food webs.
Science 279: 860–863
Pauly, D., J. Alder, E. Bennett, V. Christensen, P. Tyedmers, and R. Watson (2003). The future for fisheries.
Science 302: 1359–1361
Pew Oceans Commission (2003). America’s Living
Oceans: Charting a Course for Sea Change. Pew
Oceans Commission, Arlington, Virginia (USA)
Morgan, M.J., E.M. DeBlois, and G.A. Rose (1997). An
Pikitch, E.K.,C. Santora, E.A. Babcock, A. Bakun, R.
observation on the reaction of Atlantic cod (Gadus
Bonfil, D.O. Conover, P. Dayton, P. Doukakis, D.
morhua) in a spawning shoal to bottom trawling.
Fluharty, B. Heneman, E.D. Houde, J. Link, P.A. Liv-
Canadian Journal of Fisheries and Aquatic Science 54:
ingston, M. Mangel, M.K. McAllister, J. Pope, and
217–223
K.J. Sainsbury (2004). Ecosystem-based fishery man-
Murray, S.N., R.F. Ambrose, J.A. Bohnsack, L.W. Bots-
agement. Science 305: 346–347
MCBi.qxd
2/2/2005
326
2:41 PM
Page 326
Marine Conservation Biology
Polovina, J.J., D.R. Kobayashi, D.M. Parker, M.P. Seki,
and G.H. Balazs (2000). Turtles on the edge: Move-
Rogers, A.D. (1994). The biology of seamounts. Advances in Marine Biology 30: 305–354
ment of loggerhead turtles (Caretta caretta) along
Roman, J. and S.R. Palumbi (2003). Whales before
oceanic fronts spanning longline fishing grounds in
whaling in the North Atlantic. Science 301(5632):
the central North Pacific, 1997 – 1998. Fisheries
Oceanography 9: 71–82
508–510
Sala, E., E. Ballesteros, and R.M. Starr (2001) Rapid de-
Probert, P.K. (1999). Seamounts, sanctuaries and sus-
cline of Nassau grouper spawning aggregations in
tainability: Moving towards deep-sea conservation.
Belize: Fishery management and conservation
Aquatic Conservation: Marine and Freshwater Ecosystems 9: 601–605
needs. Fisheries 26: 23–30
Scovazzi, T. (2004). Marine protected areas on the
Ray, G.C. (1988). Ecological diversity in coastal zones
high seas: Some legal and policy considerations. In-
and oceans. Pp. 36–50 in E.O. Wilson, ed. Biodiver-
ternational Journal of Marine and Coastal Law 19(1):
sity. National Academy Press, Washington, DC
xx–xx. Available at www.iucn.org/themes/marine/
(USA)
Word/ScovazziHighSeasCOMPLETE.doc
Richer de Forges, B., J.A. Koslow, and G.C.B. Poore
Seibold, E and W.H. Berger (1993). The Sea Floor: An In-
(2000). Diversity and endemism of the benthic sea-
troduction to Marine Geology. 2nd ed. Springer Verlag,
mount fauna in the southwest Pacific. Nature 405:
944–947
Ripple, W. J. and E. J. Larsen (2000). Historic aspen recruitment, elk, and wolves in northern Yellowstone
National Park, USA. Biological Conservation 95: 361–
370
Roberts, C.M. (2002). Deep impact: The rising toll of
fishing in the deep sea. Trends in Ecology and Evolution 17: 242–245
New York (USA)
Soulé, M.E. and R.K. Noss (1998). Rewilding and biodiversity as complementary tools for continental
conservation. Wild Earth Fall, 18–28
Soulé, M.E., J.A. Estes, J. Berger, and C.M. del Rio
(2003). Ecological effectiveness: Conservation goals
for interactive species. Conservation Biology 17(5):
1238–1250
Springer, A.M., J.A. Estes, G.B. van Vliet, T.M.
Roberts, C.M. and J.P. Hawkins (1999). Extinction risk
Williams, D.F. Doak, E.M. Danner, K.A. Forney, and
in the sea. Trends in Ecology and Evolution 14(6):
B. Pfister (2003). Sequential megafaunal collapse in
241–246
the North Pacific Ocean: An ongoing legacy of in-
Roberts, C.M. and J.P. Hawkins (2000). Fully Protected
Marine Reserves: A Guide. WWF Endangered Seas
dustrial whaling? Proceedings of the National Academy of Sciences 100(21): 12223–12228
Campaign, Washington, DC (USA) and Environ-
Steele, J., ed. (2002). Effects of Trawling and Dredging on
ment Department, University of York, York (UK)
Seafloor Habitat. National Academy Press, Washing-
Roberts, C.M. and H. Sargant (2002). Fishery benefits
ton, DC (USA)
of fully protected marine reserves: Why habitat and
Terborgh J., J.A. Estes, P.C. Paquet, K. Ralls, D. Boyd-
behaviour are important. Natural Resource Modeling
Heger, B. Miller, and R. Noss (1999). Role of top car-
15: 487–507
nivores in regulating terrestrial ecosystems. Pp. 39–
Roden, G.I. (1987). Effect of seamounts and seamount
64 in M.E. Soulé and J. Terborgh, eds. Continental
chains on ocean circulation and thermohaline
Conservation: Design and Management Principles for
structure. Pp. 335–354 in B.H. Keating, P. Fryer, R.
Long-Term, Regional Conservation Networks. Island
Batiza, and G.W. Boehlert, eds., Seamounts, Islands
Press, Washington, DC (USA)
and Atolls. Geophysical Monograph, 43. American
Geophysical Union, Washington, DC (USA)
Verity, P.G., V. Smetacek, and T.M. Smayda (2002). Status, trends and the future of the marine pelagic
MCBi.qxd
2/2/2005
2:41 PM
Page 327
Place-Based Ecosystem Management in the Open Ocean
ecosystem. Environmental Conservation 29(2): 207–
237
327
Watson, R. and D. Pauly (2001). Systematic distortions
in world fisheries catch trends. Nature 414: 534–36
Walmsley, S.F. and A.T. White (2003). Influence of so-
Williams, B.K., M.D. Koneff, and D.A. Smith (1999).
cial, management and enforcement factors on the
Evaluation of waterfowl conservation under the
long-term ecological effects of marine sanctuaries.
North American Waterfowl Management Plan.
Environmental Conservation 30: 388–407
Journal of Wildlife Management 63(2): 417–440
Ward, T. and E. Hegerl (2003). Marine Protected Areas in
Wooller, R.D., J.S. Bradley, and J.P. Croxall (1992).
Ecosystem-based Management of Fisheries. Depart-
Long-term population studies of seabirds. Trends in
ment of the Environment and Heritage, Canberra
Ecology and Evolution 7: 111–114
(Australia). Available at www.deh.gov.au/coasts/
Worm, B., H.K. Lotze, and R.A. Myers (2003). Preda-
mpa/wpc/pubs/mpas-management-fisheries.pdf
tor diversity hotspots in the blue ocean. Proceedings
Warner, R. (2001). Marine protected areas beyond na-
of the National Academy of Sciences 100(17): 9884 –
tional jurisdiction: Existing legal principles and fu-
9888
ture international law framework. Pp. 55–76 in M.
Young, T.R. (2003). Developing a legal strategy for
Haward, ed. Integrated Oceans Management: Issues in
high seas marine protected areas. Pp. 81–116 in K.
Implementing Australia’s Oceans Policy. Cooperative
Gjerde and C. Breide, eds. Towards a Strategy for High
Research Centre for Antarctica and the Southern
Seas Marine Protected Areas: Proceedings of the IUCN,
Ocean, Research Report, 26. Hobart (Australia)
WCPA and WWF Experts Workshop on High Seas Ma-
Available at www.acorn-oceans.org/IOM/areas.pdf
rine Protected Areas, 15–17 January 2003, Malaga,
Watling L. and E.A. Norse (1998). Disturbance of the
Spain. IUCN, Gland (Switzerland). Available at www
seabed by mobile fishing gear: A comparison to for-
.panda.org/downloads/marine/highseamalagawork
est clearcutting. Conservation Biology 12: 1180–1197
shopproceedingsgjerdebreidehsmpa.pdf