Marine Reserves as a Tool for Ecosystem-Based Management: The
Potential Importance of Megafauna
Authors: SASCHA K. HOOKER, and LEAH R. GERBER
Source: BioScience, 54(1) : 27-39
Published By: American Institute of Biological Sciences
URL: https://doi.org/10.1641/0006-3568(2004)054[0027:MRAATF]2.0.CO;2
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Articles
Marine Reserves as a Tool for
Ecosystem-Based Management:
The Potential Importance of
Megafauna
SASCHA K. HOOKER AND LEAH R. GERBER
Marine predators attract significant attention in ocean conservation planning and are therefore often used politically to promote reserve designation. We discuss whether their ecology and life history can help provide a rigorous ecological foundation for marine reserve design. In general, we
find that reserves can benefit marine megafauna, and that megafauna can help establish target areas and boundaries for ecosystem reserves. However, the spatial nature of the interplay between potential threats and predator life histories requires careful consideration for the establishment of
effective reserves. Modeling tools such as demographic sensitivity analysis will aid in establishing protection for different life stages and distributional ranges. The need for pelagic marine reserves is becoming increasingly apparent, and it is in this venue that marine predators may be most
effectively used as indicator species of underlying prey distribution and ecosystem processes.
Keywords: marine predators, conservation, marine reserves, indicator species, modeling
The seas are by no means dead, but they are
unquestionably less alive than they were when
humanity discovered them.
—Leatherwood and Reeves (1983)
T
he state of the global oceans is rapidly deteriorating,
with dire consequences for marine species (Jackson et al.
2001). Historically, most conservation efforts have focused on
terrestrial systems, but it is becoming increasingly apparent
that conservation efforts are urgently required for the oceans
as well (Myers et al. 1997, Casey and Myers 1998). Recently,
significant attention has been given to the establishment of
marine reserves (Boersma and Parrish 1999, Mangel 2000),
with most of the focus of research directed at economically
valuable (i.e., mid–trophic level) species (Rowley 1994). Some
of the lessons learned from these reserves have now been
widely accepted (e.g., bigger is better, and dispersal matters;
NCEAS 2001). However, one of the most interesting questions
to emerge from the initial exploration of marine reserve design theory is the significance of life-history characteristics.
Here we review issues concerning the ecology of higher
predators and their relevance for the design and selection of
marine reserves.
The grouping of higher marine predators describes ocean
megafauna, including a variety of taxa: cetaceans, pinnipeds,
sea otters, polar bears, seabirds, sharks, cephalopods, and
predatory fish. Our primary expertise is in marine mammal
ecology, and so most of our review focuses on the ecology and
conservation of this group. Nevertheless, many aspects of
these species’ ecology, life history, and demography apply to
other marine predators as well, allowing us to propose certain generalities that apply to all marine predators. There is
currently a trend toward the advocation and establishment of
marine sanctuaries based on their marine megafauna, and particularly their mammal or bird fauna (table 1). However,
systematic theory on how to select, design, and monitor these
reserves is lacking, and their efficacy in protecting marine
predators is not clear. We discuss two issues here: (1) the
potential for marine reserves to protect marine predators, and
(2) the question of whether these species can serve as ecological
indicators, demonstrating where and how to target and
design marine reserves. This article is loosely based on
Sascha K. Hooker (e-mail: s.hooker@st-andrews.ac.uk) is a Royal Society
Dorothy Hodgkin Research Fellow in the Sea Mammal Research Unit at the
University of St. Andrews, Fife, KY16 9LB, Scotland. She studies marine
mammal foraging behavior, its relationship with the surrounding oceanographic
environment, and the implications for marine conservation. Leah R. Gerber
is an assistant professor in the School of Life Sciences at Arizona State
University, Tempe, AZ 85287. She works on developing approaches to connect
scientific uncertainty to decisionmaking in endangered species recovery,
marine reserve design, and disease and conservation. © 2004 American
Institute of Biological Sciences.
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Table 1. Examples of marine conservation areas established on the basis of their marine mammal and marine bird fauna.
Country/region Type of reserve
International
Europe
Year
Whale sanctuary,
1994
International Whaling Commission
European candidate special
1996
area of conservation
International sanctuary for
1999
Mediterranean cetaceans
Europe (France,
Monaco, and
Italy)
Germany
Australia
National park
Marine national park
1999
1996
Australia
Conservation park
1954
Australia
Marine park
1999
New Zealand
Mexico
Argentina
Marine mammal sanctuary
Biosphere reserve
Whale sanctuary (marine
provincial park)
Ecological reserve
National humpback whale
sanctuary
Marine national park
Pilot marine protected area
Special reservation (US
Department of the Treasury)
Fish cultural and forest
reserve (Forest Reserves Act)
Año Nuevo State Park
National marine sanctuaries
National marine sanctuary
1988
1993
1974
Brazil
Dominican
Republic
Canada
Canada
United States
United States
United States
United States
United States
Geographic area
Faunal basis for establishment
Southern Ocean
Baleen whales and sperm whale (for
recovery from historical human exploitation)
Bottlenose dolphin
Moray Firth, United Kingdom
Ligurian Sea, Mediterranean
Wadden Sea, Germany
Great Australian Bight, southern
Australia
Seal Bay, Kangaroo Island,
South Australia
Macquarie Island, Subantarctic
Fin, sperm, Cuvier’s beaked, and long-finned
pilot whales; Risso’s, striped, bottlenose,
and short-beaked common dolphins
Harbor porpoise
Southern right whale, Australian sea lion
(breeding colonies)
Australian sea lion, New Zealand fur seal
Subantarctic fur seals, Antarctic tern, fairy
prion, grey and blue petrels, and blackbrowed and wandering albatrosses (foraging
grounds)
Hector’s dolphin
Vaquita
Southern right whale
Banks Peninsula, South Island
Upper Gulf of California
Golfo San Jose,
Peninsula Valdes
Lobos Island
Silver Bank
South American sea lion and fur seal
Humpback whale
1892
Saguenay–St. Lawrence, Quebec
The Gully, eastern Canada
Pribiloff Islands (St. Paul and
St. George)
Afognak Island, Alaska
Beluga whale
Northern bottlenose whale
Northern fur seal (regulating commercial
hunt)
Seals, walrus, and sea otters
1971
1980
1992
Año Nuevo, California
Channel Islands, California
Hawaiian Islands
Northern elephant seals
Several marine mammal and bird species
Humpback whale
1983
1986
1990
1999
1869
Source: Reeves 2000.
discussion generated by a symposium that we hosted at the
annual meeting of the Society for Conservation Biology in
2001, which focused on case studies and modeling approaches
in the design of marine reserves for marine megafauna.
Marine reserves
To date, conservation work has generally employed a triage
approach: Species receive protection only after it has been
demonstrated that there is a pressing need for such protection. Many of the conservation efforts around the world,
therefore, focus on threatened or rare species (Soulé and
Orians 2001). This focus has driven much of the legislation
on conservation, which often lists species as a mechanism to
initiate efforts to protect them (see, e.g., the Endangered
Species Act and the Marine Mammal Protection Act in the
United States). However, there has been an increasing emphasis
on the need to use ecosystems, communities, and assemblages, rather than single species, as the basis for conservation.
Reserves have the potential to take this type of holistic approach (rather than traditional single-species recovery models), providing protection both to the species of concern and
to the entire ecosystem.
However, applying models developed for terrestrial systems
to the marine environment is not straightforward. Terrestrial
and marine systems are quite different ecologically in terms
of spatial structure, scale, and trophic structure (Soulé and Orians 2001). The difficulty of placing boundaries around ecosystems is exacerbated in the marine environment, where borders are dynamic and fluid. The pelagic marine environment
is vast in scale, and marine reserve areas consequently often
need to be larger than their terrestrial equivalents. There are
also differences between basic ecological structures in marine
and terrestrial environments, most notably dynamism and
connectedness (Link 2002). The spatial discreteness of terrestrial ecosystems, which allows straightforward identification of habitats to protect, is not evident in the majority of
oceanic ecosystems, which may be transient in space and
time.
Most conservation initiatives are driven by economic opportunities and constraints. In terrestrial settings, the hunting and, more recently, tourism industries have often spurred
conservation initiatives; in the marine environment, the majority of economic pressure has come from failing fisheries.
Thus, most work on marine conservation issues, and recently most evaluation of marine reserves, has been concerned with fishery recovery (NRC 2001). A recent review of
models pertaining to marine reserves (Gerber et al. 2003)
showed that none explicitly addressed reserves for top
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predators, few included explicit movement, and there was little focus on
extinction risk or multispecies interactions.
In spite of the lack of a solid theoretical foundation, large ocean
megafauna—marine mammals and
birds—are often used to direct conservation efforts (table 1, figure 1). Yet
these initiatives often have little ecological basis and are driven by public
affection toward charismatic species.
That said, marine mammals are relatively vulnerable to extinction. Of the
approximately 120 currently recognized marine mammal species, 4
species or significant populations have
gone extinct, 11 are thought to be in
imminent peril of extinction, 17 are
thought to be of significant concern
with respect to extinction, and 8 were
once thought to be at risk of extinction
but are now recovering (table 2; VanBlaricom et al. 2000).
a
b
Definitions and goals
We define a marine protected area as
a geographic area designated for protection. This may include a broad area
with limited management restrictions
(e.g., prohibiting some activities such
as seismic exploration) but may also
encompass smaller “marine reserve
areas”—zones designated as closed to
extraction (NRC 2001). In this article, we focus on the degree to which the
spatial nature of marine protected
areas can promote recovery and en- Figure 1. (a) Bottlenose whales in the Gully, eastern Canada. The Gully has been
hance protection from the threats that designated a pilot marine protected area, largely because of the northern bottlenose
marine predators face. The question of whales found there. These whales often spend periods of time resting at the surface
whether marine reserves will provide between foraging dives. Threats to these and other cetaceans in this region include
protection and will prohibit the activi- ship strikes, noise pollution from exploitation and exploration, and interactions with
ties that threaten these predators longline fisheries. Photograph: Hal Whitehead laboratory. (b) Humpback whales in
or their ecosystems is a key issue. A Hawaii. The Hawaiian Islands Humpback Whale National Marine Sanctuary was
major criticism of marine reserves gen- established to protect breeding humpback whales. In general, threats are low in this
erally, and particularly several of those area, although a growing whale-watching industry and acoustic testing nearby may
established for marine mammals, is be causes for concern. Photograph: Robin W. Baird.
that they represent “paper parks” that
provide a false sense of conservation achievement (Duffus and
harvested fish or invertebrate populations to help support
Dearden 1995). This criticism stems from the lack of regulafisheries outside the reserve. The ranking of these goals will
tion and policing or wardening for such reserves or sanctudepend on the societal and economic pressures for a given
aries (Hooker et al. 1999).
region. Here, for the most part, we focus on the impact of
The goals of establishing a marine protected area are sevreserves on higher predators, although we also consider
eral: conservation of biodiversity (minimizing extinction
multispecies and multipurpose reserves in terms of whether
risk), ecosystem protection, reestablishment of ecosystem
fishery enhancement is possible in conjunction with conserintegrity, and enhancement of the size and productivity of
vation of higher predators.
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Table 2. Marine mammal species, subspecies, and populations that are extinct, at risk of extinction, or recovering from
near-extinct status.
Status
Species (population)
Latin name
Extinct (4)
Steller’s sea cow
Caribbean monk seal
Japanese sea lion
Gray whale (North Atlantic)
Hydrodamalis gigas
Monachus tropicalis
Zalophus japonicus
Eschrichtius robustus
In imminent peril of extinction (11)
Baiji
Vaquita
Indian river dolphin (Indus river)
Mediterranean monk seal
Gray whale (western North Pacific)
Right whale (eastern North Pacific)
Right whale (North Atlantic)
Bowhead whale (Davis Strait, Hudson Bay,
Spitsbergen, Barents Sea, and Sea of Okhotsk)
Beluga whale (Gulf of Alaska)
Beluga whale (Gulf of St. Lawrence)
Ringed seal (Lake Saimaa)
Lipotes vexillifer
Phocoena sinus
Platanista gangetica
Monachus monachus
Eschrichtius robustus
Balaena glacialis
Balaena glacialis
Balaena mysticetus
Of significant concern (17)
Blue whale
Hawaiian monk seal
Ringed seal (Baltic Sea)
Ringed seal (Lake Ladoga)
Harbor seal (western North Pacific)
Steller’s sea lion (western North Pacific)
Australian sea lion
Hooker’s sea lion
Guadalupe fur seal
Juan Fernandez fur seal
Walrus (Atlantic)
Walrus (Laptev Sea)
Amazonian manatee
West African manatee
West Indian manatee
Sea otter (California)
Marine otter
Balaenoptera musculus
Monachus schauinslandi
Pusa hispida botnica
Pusa hispida ladogensis
Phoca vitulina stejnegeri
Eumetopias jubatus
Neophoca cinerea
Phocarctos hookeri
Arctocephalus townsendi
Arctocephalus philippii
Odobenus rosmarus rosmarus
Odobenus rosmarus laptevi
Trichechus inunguis
Trichechus senegalensis
Trichechus manatus
Enhydra lutris nereis
Lutra felina
Once thought to be faced with
extinction but now recovering (8)
Bowhead whale (western Arctic)
Humpback whale
Gray whale (eastern North Pacific)
Northern elephant seal
Galapagos fur seal
Subantarctic fur seal
Antarctic fur seal
Sea otter (North Pacific and Russian coastal waters)
Balaena mysticetus
Megaptera novaeangliae
Eschrichtius robustus
Mirounga angustirostris
Arctocephalus galapagoensis
Arctocephalus tropicalis
Arctocephalus gazella
Enhydra lutris kenyoni
Delphinapterus leucas
Delphinapterus leucas
Pusa hispida saimensis
Source: VanBlaricom et al. 2000.
Marine predators and marine reserves
Threats to marine predators may take several forms (box 1,
figure 2; Richardson et al. 1995, Simmonds and Hutchinson
1996, Coe and Rogers 1997). Physical threats may include
strikes from ships or entanglement by fisheries, often leading
to the death of individual animals. Acoustic or environmental impacts may be more insidious. Seismic exploration, military exercises, shipping, or drilling may have far-reaching
acoustic impacts that cause species to leave an area, to become
temporarily unable to forage, or even to sustain physical
damage. Similarly, pollution, dumping, and oil spills may
increase the risk of extinction by increasing mortality. Ingestion
of plastic debris, oil contamination, and pollutants may have
an incremental effect on animals throughout their lives,
ultimately resulting in immunosuppression or reproductive
failure. Potentially irreparable ecosystem changes caused
by competition for resources may radically alter ecosystem
structure, resulting in dramatic shifts in population demographics (e.g., the Southern Ocean; May 1979), and habitat
disturbance or destruction can result in spatial shifts to
distribution or migration routes due to loss of cultural
memory. Many of these threats may be mitigated by spatial
protection.
In protecting a specific population, the optimal protected
area would encompass that population’s year-round distribution (Reeves 2000). However, for many marine predators, the
year-round distribution of a population may span entire
ocean basins. The question therefore becomes whether limited spatial protection in specific parts of a species’ range is
worthwhile. In some cases, when only a portion of a wideranging predator population may use a protected area, there
may be the potential for recolonization of overexploited
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Box 1. Threats to higher predators
and to the ecosystem
Direct threats
Direct threats are those that cause mortality of top predators.
Fishery bycatch. Several seabird and cetacean species are
killed in fisheries around the world. The establishment of
reserves can mitigate these population-level impacts and
reduce exposure at an individual level.
Direct killing. In some places, seabird, cetacean, and pinniped
species are still the focus of directed hunts.
Ship strikes. In certain areas there are increased risks of ship
strikes. For instance, in the Bay of Fundy, in the northeastern
United States, ship traffic en route to Boston presents a large
threat to northern right whales.
Indirect threats
Rather than causing immediate death, these insidious threats
may cause accumulating harm over longer time scales.
Overexploitation of lower trophic levels. By removing lower
trophic levels from the food chain, nutritional stress may be
imposed on upper trophic predators, causing switching of
prey, offspring desertion, and, in extreme cases, starvation.
Habitat degradation. This may take a variety of forms:
•
Acoustic pollution. This can cause potential immediate
damage to soft tissues in the case of high-intensity
sounds, or behavioral avoidance of an area in the case of
lower-intensity sounds.
•
Chemical pollution. This can affect all levels of the food
chain but becomes bioaccumulated at increasing trophic
levels. Among higher predators, it can cause immunosuppression and potential reproductive failure.
•
Marine debris. Animals may become entangled in discarded rope and nets, or materials such as plastic bags
may be ingested.
•
Physical habitat destruction. Some trawling methods
cause long-lasting damage to the sea floor, which may
take decades to recover to its previous physical structure.
Similarly, because pinnipeds and seabirds rely on terrestrial sites for breeding, they may also be susceptible to
habitat degradation and disturbance (e.g., invasive vegetation, introduced predators, light pollution).
Global effects
Global effects, such as climate change, will have consequences
for higher predators and their marine ecosystems. These
threats require mitigation at a global level.
subpopulations because of migration from this viable subpopulation (i.e., source–sink dynamics). However, even when
this is not the case, the establishment of areas in which these
threats are reduced or removed can only be beneficial, since
several of the threats faced by marine mammals are site specific and others have cumulative effects (box 1). Even if a
predator used the protected area for only a portion of its life
span, this would reduce the frequency with which each individual was exposed to certain impacts and diminish the overall cumulative impact of other threats.
Modeling approaches can be usefully applied to the question of when and how to protect different life stages and distributional ranges to promote population protection. The annual cycle of many higher predators consists of discrete
foraging and breeding portions. Thus, both foraging and
breeding habitat and the migratory route should be considered (figure 3). Demographic rates may differ with the annual
cycle or with specific habitats, and consideration of these
variations may help to prioritize potential reserve sites. Similarly, the vulnerability of the population may be habitat and
stage specific. For example, evidence of depressed breeding
success due to local food limitation or disturbance at breeding sites would suggest that enhancement of the breeding population (e.g., by enhancing reproductive success or breeding
population size) would be of most value. Thus, when this is
the case, reserves should be established around breeding
areas to protect important food resources during the breeding season along with the breeding individuals themselves.
Conversely, evidence of food limitation or population-level
threats during foraging would suggest that protected areas
need to be established at sea, away from breeding colonies.
Likewise, if migrating individuals become spatially concentrated at particular ocean areas or near particular features,
where they could be especially vulnerable to threats such as
ship strikes or bycatch, then establishing reserves in these
areas would be warranted.
The current criteria used to select reserves for marine
mammals have primarily involved the identification of breeding areas and have only occasionally taken account of foraging or migration habitats (table 1). Among pinnipeds, seabirds,
and turtles, breeding and pupping or nesting takes place at
terrestrial sites, where breeding animals are frequently highly
aggregated. Part of the impetus for selection of such sites is
likely to have come from their ability to encompass high
spatial aggregations of individuals in a relatively small protected area, rather than from a thorough consideration of
spatial and demographic threats. Future research should
couple quantitative approaches to predict reserve efficacy
with field studies of habitat use (i.e., the importance of foraging, migration, and breeding habitats). On a global scale, an
examination of the threats faced by these animals (box 1) suggests that it is likely to be during foraging that most individuals are at risk, and it is here that research attention needs to
be directed.
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a
b
d
c
e
Figure 2. Threats to marine mammal species may be direct, including (a) fishery bycatch (harbor porpoise caught accidentally in a fishery; photograph: Nigel Godden, Sea Mammal Research Unit) or (b) ship strikes (propeller wound in right
whale; photograph: Robin W. Baird), or indirect, including (c) debris (Antarctic fur seal entangled in discarded fishing gear;
photograph: Sascha Hooker), (d) reduction in prey because of fisheries (photograph: Dave Sanderson, Sea Mammal
Research Unit), or (e) oil and gas activity (pilot whales swimming beside rig, eastern Canada; photograph: Robin W. Baird).
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Modeling approaches and
the influence of life history
from annual (in the case of migratory species) to decadal (in
the case of species following El Niño effects). Most of the models that have incorporated animal movement have taken little of this into account (Roberts and Sargant 2002). Few existing models have considered all life stages; thus, most have
failed to acknowledge that wide-ranging marine species may
have life-history stages that occur in very different habitats.
Much of the theory developed for marine reserves has instead
focused on issues of larval dispersal; very few studies have addressed the question, “For which types of species should
reserves be most effective?” Demographic population
models (e.g., Caswell 2001) are one promising approach to
examining the potential efficacy of marine reserves that
target particular life-history stages or their habitats.
To date, marine reserves have been developed with little
scientific basis to assess the effectiveness of various reserve
designs and with few quantitative approaches to monitor
these reserves. Models of marine reserves are relatively new
in the literature; Gerber and colleagues (2003) reviewed 34
articles concerning marine reserve modeling, of which 32 were
published after 1990. Based on these existing models of
marine reserves, they reported that reserves will provide
fewer benefits for species with greater adult rates of movement.
However, few models have been developed explicitly for
wide-ranging species. Those studies that have included
migration and movement in reserve models show continued
benefits even to highly mobile species
(Apostolaki et al. 2002, Roberts and
Sargant 2002); there is also empirical support for this (Gell and Roberts 2003).
While models of marine reserves are bem1
ginning to yield information on the necessary spatial configurations of reserves to
allow populations with specific dispersal
m2
distances to persist, spatial configuration
remains an aspect of reserve design in need
of further analysis. Important directions
for future modeling include the effects of
1 – m1
particular forms of density dependence and
multispecies interactions and the consideration of full life-stage models in reserve
design. This additional modeling and analy1 – m2
sis will improve prospects for a better understanding of the potential of marine
reserves for conserving biodiversity.
The major difference between the modeling approaches we advocate for higher
predators and those previously used for
fisheries involves the explicit consideration Figure 3. Life stages of some marine predators (e.g., baleen whales, pinnipeds,
of life-history strategies, with conservation seabirds) are separated spatially into discrete feeding and breeding areas, with
goals operating over much larger spatial migration between them. Reserves can be placed in feeding, breeding, or migraand temporal scales. The multispecies tory habitats. Abbreviations: m, migration rate (m and m indicate different
1
2
nature of most marine ecosystems also ne- rates for migration to each feeding area); S, mixing between feeding areas.
cessitates delicate structuring of conservation priorities between ecosystem levels. We propose that
Demographic sensitivity analysis allows researchers to
modeling tools such as demographic sensitivity analysis
analyze how much a small change in a demographic rate
and multispecies models may be worth consideration to
(e.g., adult survival) would influence a population’s potential
explore approaches for the conservation of higher marine
for recovery. Further, such approaches rely on minimal data
predators.
(e.g., survival and fecundity rates) and may allow researchers
One interesting facet of the debate on reserve design is that
to predict the effects of various management actions. Altermost reserves are quite small in area and represent only a small
native designs for marine protected areas should be treated
portion of the total range of species. At the same time, modelas hypotheses and tested with models in advance (Heppell and
ing work suggests that as movement rates increase, larger
Crowder 1998). Because life-history information is lacking for
areas are needed for reserves to achieve benefits (Gell and
many marine populations, categorizing life histories accordRoberts 2003, Gerber et al. 2003). Whereas fish may dising to their response to changes in stage-specific mortality may
perse, and therefore an individual fish may move away from
provide a useful framework for considering conservation
an area for the majority of its life, many higher predator
options (Heppell et al. 2000a). For example, it is well estabspecies are relatively site faithful over time scales varying
lished that fecundity will be more important than survival
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for shorter-lived species, such as many fish and invertebrates,
whereas adult and juvenile survival elasticity will be important for long-lived species such as marine mammals, sea turtles, and seabirds (Heppell et al. 2000b, Saether and Bakke
2000).
These insights from life-history theory can be used to predict a priori when marine reserves are likely to be most effective, and perturbation analysis can serve as an early step
in reserve planning. Perturbation analysis is useful to show
that different life-history characteristics will exhibit variable
responses to changes in mortality. For example, across a
representative range of marine life histories (e.g., urchin,
haddock, sea lion), the change in population growth rate (λ)
resulting from a decrease in adult mortality will be greatest
for marine invertebrates and fish, and lowest for species
with very low adult mortality rates (e.g., salmon will show
a more striking response than sea lions). Results from demographic analysis suggest that adult survival rate and
maximum life span are critically important in determining
reserve efficacy. Standardized demographic analysis (sensu
Caswell 2001) may be a useful first step to compare disparate
conservation goals for marine reserve design for species
with distinct life histories.
Multispecies and habitat-based models
Predictions of the effect of a reserve on a particular species may
also vary depending on species interactions. Thus, a major
challenge in marine reserve design is the incorporation of
multispecies interactions and management objectives. Decades
of experimental studies have shown dramatic effects of
consumer–resource interactions on populations and communities (Soulé and Orians 2001). The relationship between
marine predator distribution and areas of primary productivity has been observed for several species (McConnell et al.
1992, Jaquet and Whitehead 1996). Bottom-up effects of resources on consumers and top-down effects of consumers on
other species in the community often include a suite of direct
and indirect pathways of interaction (Bowen 1997). Predator–prey, competitive, and mutualistic interactions can cause
unanticipated changes in community structure and nontarget effects of management interventions. Adding species interactions to predator–prey models to explore the effects of
different reserve designs can generate complex responses to
protection, changing not only the magnitude but also the direction of the species response. Previous perturbations and
reduction in certain components of a food web relative to
other components mean that management actions to restore
the system may result in an oscillation that causes unforeseen
consequences and, in the worst case, a complete ecosystem shift
(Estes et al. 1998). In addition, population size and ecosystem
effects may be linked with disease outbreak. In general, maintaining high trophic connectivity and preventing competitive
release that leads to abnormally elevated population levels will
decrease the levels and impact of disease (Lafferty and Gerber 2002).
Multipurpose reserves
Can enhancement of fisheries (e.g., increase in the size and
productivity of harvested fish and invertebrate populations
to help support fisheries outside reserves) be viable alongside
reserves that maintain higher predator numbers? In California, marine reserves have been established to protect depleted and reintroduced sea otters. However, there is also an
interest in the promotion of abalone populations, which
have expanded in the absence of sea otters. Sea otters in
Alaska have been shown to exert a profound effect on the
structure of their marine community, encouraging kelp
growth through predation on slow-moving herbivorous
invertebrates (Estes and Palmisano 1974). Such cascading
ecosystems are governed by the strength of the trophic link
between otters and invertebrates relative to other factors,
such as recruitment variability or natural disturbance, that
affect biodiversity in kelp forests. With strong linkage, the presence of sea otters can provide a control on the herbivore
population, enhancing the overall biodiversity of kelp forest
ecosystems, increasing the amount of productivity and pathways through the food web, and promoting structural complexity in the ecosystem (Estes et al. 1998). In such cases, it
would appear that sea otter presence within marine reserves
is desirable.
However, recent empirical work has shown that the strength
of sea otter predation on abalone was greater than the pressure of the fishery on abalone, so that there were fewer abalone
in reserve areas containing sea otters than in fished areas
with no sea otters present (Fanshawe et al. 2003). Thus, reserves
that maintain ecosystem integrity and natural (unperturbed)
levels of predation do not appear to be consistent with additional abalone harvesting. While this example suggests that
there may be situations in which marine reserves cannot
simultaneously protect multiple trophic levels, one approach
to addressing this particular issue is to spatially separate
areas into two single-use marine protected areas: one focusing on restoring ecosystem integrity and the other focusing
on protecting the harvested stocks to enhance productivity
(Fanshawe et al. 2003).
This conflict, like many conflicts between fisheries and
conservation of higher predators, has arisen largely because
of historical changes in the ecosystem (Jackson et al. 2001).
Removal of sea otters caused increases in the population
sizes of lower levels of the food web. Fanshawe and colleagues
(2003) suggest that an incomplete ecosystem may provide
greater value to human consumers than the restored system.
However, in general, it is thought that increased food web complexity leads to greater ecosystem resilience, and a restored
ecosystem would be preferred on this basis (Soulé and Orians 2001). The value of the fishery economically and sociologically must therefore be weighed against the risks associated with maintenance of the perturbed ecosystem.
Marine predators as indicator species
In terrestrial ecosystems, predator distribution can be used to
establish criteria for reserve design (Soulé and Simberloff
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1986), but the application of this concept to oceanic systems
is relatively untested. Larger predators have been used as
indicator (or focal) species, whose protection aids in protecting
the more complex environments that they use (Simberloff
1998, Zacharias and Roff 2001). There are several variants on
the meaning of focal species, and marine predators have been
used to address most of these concepts (table 3). In general,
although these concepts may be useful to drive conservation
efforts, the majority are focused on achieving political support through publicity or are extensions of single-species
conservation efforts (Simberloff 1998, Andelman and
Fagan 2000). Of all these, the indicator-species approach
appears to have the most potential to direct marine reserve
selection (Zacharias and Roff 2001).
Another related approach is to identify biodiversity hotspots
that are worthy of protection. These may be areas that are particularly rich in species, in rare species, in threatened species,
or in some combination of these attributes (Reid 1998,
Myers et al. 2000). In the marine environment, Roberts and
colleagues (2002) found that centers of endemism among coral
reef systems are major biodiversity hotspots. However, there
is still much disagreement in the academic community over
what constitutes biodiversity. Is species richness the answer,
or are these simply locations in which several species overlap,
possibly at the edges of their ranges (Price 2002)? We would
suggest that, at least in the pelagic realm, a more appropriate
definition of oceanic hotspots may be areas of increased productivity, in terms of the abundance of organisms within
the area relative to other oceanic areas. The Gully, a submarine canyon offshore of eastern Canada, appears to be a
hotspot for cetaceans, which show elevated abundances in the
vicinity of this feature compared with the levels in surrounding regions (Hooker et al. 1999). Similarly, the Patagonian shelf has been demonstrated to be a rich feeding
ground for several predator species in the south Atlantic
(Croxall and Wood 2002).
There is some evidence that the distribution and relative
abundance of marine predators can be used as an indication
of underlying prey distributions and ecosystem processes
(Preen 1988, Tershy et al. 1991, Croll et al. 1998). In the Gully,
although the underlying process driving this ecosystem is
not well understood, top predators could be used to help
derive boundaries for protection (Hooker et al. 1999, 2002).
Although this area is relatively far offshore, the distribution
of cetaceans is primarily governed by bathymetric features and
so could be well defined by spatial boundaries. Such associations have not been found for other areas, and offshore
environments are generally more dynamic, making it difficult
to establish distinct spatial boundaries. However, establishing such boundaries may be a matter of directing research
attention to oceanographic features that may show stability
in space and time. Thus far, conservation in the pelagic realm
has received little attention, although there have been suggestions that a system of open ocean reserves is needed (Mills
and Carlton 1998).
The identification of foraging hotspots for predators (see
figure 3) and the consideration of boundaries determined by
oceanographic processes show potential as useful approaches
in the pelagic arena. Hyrenbach and colleagues (2000) identified three types of oceanic hotspots: (1) static systems,
which are determined by topographic features; (2) persistent
hydrographic features, such as currents and frontal systems;
and (3) ephemeral habitats, shaped by wind- or currentdriven upwelling, eddies, and filaments. Of these, ephemeral
habitats are by far the most difficult to map or protect. All three
types of hotspots may be identified by analyzing the foraging distribution of higher predators (table 4). Thus, the overlaying maps of different marine predators’ foraging habits,
together with basic knowledge of their diet (e.g., piscivory or
teuthophagy), broader ecosystems (e.g., upwelling dynamics),
and habitat variability (e.g., persistence and spatial variation
over annual and decadal cycles), should allow researchers to
identify various hotspot features.
Networks of marine reserves
The choice of new areas to protect will necessarily be influenced by the types of areas that are already being protected.
A good deal of research effort has been expended in assess-
Table 3. Focal species concepts, with examples from marine mammal fauna.
Type
Description
Example
Flagship
Charismatic species that serve as guarantors of broadscale conservation, used politically to attract funding
and support
Right whales or humpback whales (national
marine sanctuaries, United States)
Keystone
Species that have a disproportionate effect relative to their
abundance, underpinning the ecosystem
Sea otters
Umbrella
Wide-ranging species, the protection of whose habitat will
encompass several other species within their ecosystem
Manatees, river dolphins
Composition indicators
Species whose presence or abundance is used to
characterize a particular habitat or biological community
Northern bottlenose whales (pilot marine
protected area, the Gully, Canada)
Condition indicators
Species that reflect ecosystem health or the levels of
pollutants within the system
Arctic cetaceans
Source: Simberloff 1998, Zacharias and Roff 2001.
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Table 4. Types of oceanic hotspots and examples of top predator distribution associated with them.
Type of hotspot
Location
Higher predator
Static systems
The Gully, Nova Scotia
Northern bottlenose distribution is driven by the bathymetry of
the submarine canyon (Hooker et al. 1999).
Hawaiian Islands
National Marine Sanctuary
Humpback whale distribution in the breeding season is
primarily defined by the shallow water region around the
Hawaiian Islands.
Persistent hydrographic features
Bird Island, South Georgia
Antarctic fur seal and macaroni penguin distribution is found to
the northwest of South Georgia, where there appears to be a
persistent frontal system (Barlow et al. 2002).
Ephemeral habitats
Warm core ring, North Atlantic
Sperm whales are found primarily along the periphery of warmcore rings (Griffin 1999).
Note: Hotspot type is derived from Hyrenbach and colleagues (2000).
ing optimality in networks of protected areas. Explicit, quantitative methods of identifying priority areas for biodiversity
are replacing the ad hoc procedures often used in the past to
design networks of reserves. The concept of complementarity ensures that areas chosen for inclusion in a reserve network
complement those already selected, reducing duplication of
species in reserves and providing the most efficient network
of protected areas. The network of reserves is identified based
on uncorrelated habitat types or assemblages to provide a network of protected areas encompassing a high proportion of
biodiversity. This concept is gaining support in terrestrial
reserve network assessment (Howard et al. 1998, Reyers et al.
2000), but it could be applied equally well to marine systems.
Furthermore, although in the past this method has relied on
high-quality information on the spatial distribution of all
species of concern, it appears that reserve selection based on
data obtained with low sampling effort can be highly effective in the representation of species (Gaston and Rodrigues
2003). This is likely to have important consequences in the
identification of marine mammal habitats, which are generally poorly known, but for which peaks of abundance are better documented.
Socioeconomic concerns also strongly influence the decisionmaking process for the establishment of networks of reserves. In the southern Gulf of California, multiple levels of
information on biodiversity, ecological processes, and socioeconomic factors were used to establish a network of reserves that would cover a large proportion of habitat and reduce social conflict (Sala et al. 2002). A major benefit of the
optimization technique used in creating this network is the
ability to generate portfolios of solutions that can be presented
to decisionmakers, who can evaluate the costs and benefits of
different management options within relevant socioeconomic constraints.
Management case studies
Two political vehicles for protected area management are
illustrated in boxes 2 and 3. In the European Union, a recent
directive (Council Directive 92/43/EEC) has led to the need
to establish special areas of conservation for species and habi-
tats (box 2). Five marine mammal species require protection
under this scheme. The ease of designating these areas reflects
many of the issues we have discussed here. The protection of
two sites encompassing large colonies of breeding grey seals
and the protection of small, localized bottlenose dolphin
populations have been fairly straightforward. However, more
broadly distributed species without large breeding aggregations are more difficult to assess. For instance, harbor porpoises
do not appear to show site fidelity, and the relationship between their foraging areas and local oceanography is unknown. Setting up a marine reserve for harbor porpoises
would therefore require the zoning of large offshore areas, a
prospect that is unlikely to be politically or economically
palatable.
At a much larger scale, the Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR) has
been established to protect the Southern Ocean (box 3).
CCAMLR is heavily management driven, concerning itself primarily with catch quotas and fishery licensing. Nevertheless,
one of its mandates is to ensure that the other ecosystem
components of these fisheries are not adversely affected.
Thus, the survival and growth rates of higher predators (pinnipeds and seabirds) are monitored and the fishery adjusted
accordingly (Constable et al. 2000). Thus far, this legislation
has not considered the possibility of using offshore reserves,
although concerns over seabird bycatch may encourage such
an approach in the near future.
Monitoring and policing
One of the greatest limitations facing novel conservation initiatives is the difficulty involved in assessing their effectiveness
at protecting target species. Once a reserve area or management strategy has been implemented, can researchers assess
whether it has slowed the rate of decline of the target species?
Most higher predators have long life spans, and consequently
it is often several years before any changes in population
growth or structure become apparent. A potentially promising method to investigate this is to look at shifts in age structure. Although these shifts will also take several years to observe, there is usually a transitory oscillation immediately
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Box 2. Case study: The European Union’s
special areas of conservation
The recent European Council Directive 92/43/EEC, also known as
the Habitats Directive, was passed to provide protection to 632
species and 56 habitat types within Europe. This legislation
requires European countries to designate special areas of conservation (SACs). There are three main criteria for the designation of
these areas:
• Each area must contain priority species that are rare in the
country where the area is located.
• If possible, an area should be chosen to protect both specific
species and specific habitats, not simply by drawing a box
around species distribution but by identifying and incorporating habitat types around a species distribution.
• Within the European Union (EU), a high proportion of the
entire population of the priority species should be conserved.
Fifty to ninety percent of the EU populations of most marine
mammals are found within the United Kingdom (UK), placing a
requirement on this country to develop plans for designating and
implementing reserves for these species.
Five marine mammal species require protection:
• Grey seal (UK population approximately 130,000). Breeds
in localized colonies on islands around Scotland. Primary
threat is to breeding rather than foraging sites. Three to four
SACs will encompass approximately 50% of the breeding
population.
• Bottlenose dolphin (UK population approximately 300–500).
Localized areas of distribution in the Moray Firth, Scotland,
and Cardigan Bay, Wales. Each SAC encompasses a large
portion of the distribution of a local population.
• Harbor seal (UK population approximately 50,000). Hauls
out in diffuse groups at several locations; no large colonies.
High haul-out densities along the west coast of Scotland,
Shetland and Orkney, but would require a number of different sites to protect a significant portion of the population.
Conflicts with inshore fishermen. Also vulnerable to occasional epidemics.
• Mediterranean monk seal (European population approximately 500). Population has suffered catastrophic collapses,
leaving remnant groups of animals. Vulnerable to disturbance at haul-out and breeding sites. Protection of viable
breeding habitat is encouraging recolonization; however, 2
areas in Portugal and 11 in Greece are probably protecting
fewer than 100 individuals.
• Harbor porpoises (North Sea population approximately
340,000). Areas of high density offshore are dispersed and
mobile; a vast protected area would be required to protect
survival and reproduction. This has led to a stalemate in
terms of reserve designation.
after the successful implementation of a management
action, which may be observed using proxies for age structure
(e.g., juvenile-to-adult ratios; Holmes and York 2003).
In addition to monitoring the progress of protected populations in the context of management goals (e.g., recovery
from overexploitation), it is important to document levels of
illegal take from hunting or fishing in protected areas. Unless
there is strong community support for a particular marine
protected area, there is likely to be some take within the area
that may obscure detection of its protective effects or undermine those effects completely. Similarly, the policing of marine protected areas is extremely difficult, particularly within
the highly dynamic pelagic system. The pirate fishery for
Patagonian toothfish in the Southern Ocean is an example of
this, occurring within the CCAMLR management system
(Constable et al. 2000). This is where economics may provide
the simplest answer—it will only be by controlling the
market that such pirate fishing may become unsustainable.
Conclusion and future directions for research
Can new management tools such as marine reserves be useful for conserving marine megafauna? Although marine
mammals and birds have traditionally been used as flagship
species for conservation efforts, novel designs of marine protected areas guided by a consideration of these species’ distribution and life history may greatly enhance the effectiveness of existing protective measures. We have discussed several
issues that will play a role in developing such designs. Most
important, assessment of the threats that will be mitigated by
a reserve, and consideration of the anticipated management
actions, should be incorporated at an early stage. Consideration of such threats in combination with distribution and
life-history data will help to establish the size and placement
of protected areas. In terms of modeling options, demographic sensitivity analysis may be relevant to the question of
when and how to protect different life stages and distributional
ranges to promote population protection. In particular,
alternative designs for marine protected areas can be compared
with demographic and ecological models before experimental reserves are established (Heppell and Crowder 1998).
The impacts of reserves on species other than their targets
are more difficult to predict, and future work is needed on the
dynamics of interspecific interactions associated with the
recovery of populations. However, in systems that have not
suffered large-scale ecological perturbation, or systems in
which reserves will serve to provide precautionary protection,
this will not be an issue.
The use of marine predators as indicator species may provide a useful approach to the protection of productive ocean
areas. Existing knowledge of areas that represent peaks of
abundance for marine megafauna should enable establishment
of networks of pelagic reserves based on distributional
hotspots and complementary species protection (Reyers et al.
2000, Gaston and Rodrigues 2003). The final limiting factor
for such reserves is more likely to be the political will and
international cooperation necessary to achieve them.
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Box 3. The Southern Ocean: Convention for the
Conservation of Antarctic Marine Living Resources
In contrast to other multilateral fisheries conventions, the
Convention for the Conservation of Antarctic Marine Living
Resources (CCAMLR; www.ccamlr.org) not only is concerned
with the regulation of fishing but also has a mandate to conserve the entire Antarctic marine ecosystem. CCAMLR was a
pioneer in developing this ecosystem approach to the regulation of fisheries, considering the whole Southern Ocean as a
suite of interlinked systems.
A conventional definition of an ecosystem is any unit that
includes all of the organisms in a given area, interacting with
the physical environment so that a flow of energy leads to
clearly defined trophic structures, biotic diversity, and material cycles (i.e., exchange of materials between living and nonliving parts) within the system. An ecosystem approach does
not concentrate solely on the species fished; it also seeks to
minimize the risk that fisheries will adversely affect “dependent and related species,” that is, species with which humans
compete for food. However, regulating large and complex
marine ecosystems is a task for which managers currently
have neither sufficient knowledge nor adequate tools. Instead,
CCAMLR’s approach is to regulate human activities (e.g.,
fishing) proactively so that deleterious changes in the Antarctic ecosystems are avoided.
CCAMLR applies to all areas south of 60° S and to waters
between that latitude and the Antarctic Convergence, the
oceanographic boundary between the Southern Ocean and
other global oceans. Its goals are
• to facilitate research into and comprehensive studies of
Antarctic marine living resources and the Antarctic
marine ecosystem
• to compile data on the status of and changes in populations of Antarctic marine living resources
• to ensure the acquisition of catch-and-effort statistics
on harvested populations
• to identify conservation needs and analyze the effectiveness of conservation measures
However, the use of marine reserves for conserving marine
megafauna will be of limited value without the backup of firm
management guidelines. Scientists and managers need to become less accepting of having areas designated as sanctuaries without tangible protection. Of course, with any management restrictions, enforcement will be a problem,
particularly on the high seas. For effective conservation, a
change in public sentiment may be required, such that ocean
users become more effective at policing themselves. Ultimately, conservation benefits in the ocean are likely to depend
on greater vision on the part of scientists—and, most critically, of policymakers—in realizing the benefits of favoring
long-term sustainability over short-term economic profit.
Acknowledgments
Many thanks to symposium participants Sarah Allen, Dee
Boersma, Ian Boyd, Andy Dobson, John Harwood, Eli Holmes,
David Hyrenbach, Kevin Lafferty, Fiorenza Micheli, and
Glenn VanBlaricom for interesting and thought-provoking
presentations and discussion. The manuscript benefited from
review by Ian L. Boyd, Paul K. Dayton, David Hyrenbach,
Simon Northridge, Callum M. Roberts, Glenn VanBlaricom,
and one anonymous reviewer.
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THE BEST IN BIOLOGICAL SCIENCES FROM SPRINGER!
COMPUTATIONAL CELL BIOLOGY
CHRISTOPHER FALL, New York University, New York, NY;
ERIC MARLAND, Appalachian State University, Boone, NC;
JOHN WAGNER, University of Connecticut Health Center, Farmington, CT;
and JOHN TYSON, Virginia Polytechnic Institute & State University,
Blacksburg, VA (Eds.)
This textbook provides an introduction to
dynamic modeling in cell biology, emphasizing computational approaches based on
realistic molecular mechanisms. It is designed
to introduce cell biology and neuroscience
students to computational modeling, and
applied mathematics students, theoretical
biologists, and engineers to many of the
problems in dynamical cell biology. Illustrative
exercises are included with every chapter, and
mathematical and computational appendices
are provided for reference. This textbook
will be useful for advanced undergraduate and graduate theoretical
biologists, and for mathematics students and life scientists who wish
to learn about modeling in cell biology.
Royalties from this book will be donated to the Joel E. Keizer memorial
endowment for collaborative interdisciplinary research in the life
sciences.
2002/488 PP., 210 ILLUS./HARDCOVER/$59.95
ISBN 0-387-95369-8
INTERDISCIPLINARY APPLIED MATHEMATICS, VOL. 20
INSTANT NOTES IN MOLECULAR BIOLOGY
Second Edition
P.C. TURNER, A.G. MCLENNAN, A.D. BATES, and M.R.H. WHITE, all,
University of Liverpool, UK
Instant Notes in Molecular Biology provides
a structured approach to learning by covering
all the important topics in a uniform, systematic
format. Each topic begins with a summary of
the essential facts — an ideal revision checklist — followed by detailed explanation and
clear, simple diagrams. The diagrams are
particularly easy to learn and reproduce for
essays and examinations. Since their launch
in 1997, the Instant Notes books have become
bestsellers and this new edition, which has
been completely updated, will remain very
attractive to any undergraduate student taking
a course on molecular biology.
2000/360 PP., 160 ILLUS./SOFTCOVER/$29.95
ISBN 0-387-91601-6
INSTANT NOTES
Journals
ECOSYSTEMS
PRINCIPLES OF TERRESTRIAL
ECOSYSTEM ECOLOGY
F. STUART CHAPIN III, University of Alaska, Fairbanks, AK;
PAMELA MATSON and HAROLD A. MOONEY, both, Stanford
University, Stanford, CA
The ecosystem approach to ecology treats
organisms and the physical elements of
their environment as components of a single,
integrated system. This comprehensive
textbook outlines the central processes that
characterize terrestrial ecosystems, tracing the
flow of water, carbon, and nutrients from their
abiotic origins to their cycles through plants,
animals, and decomposer organisms. This
book synthesizes current advances in ecology
with established theory to offer a complete
survey of ecosystem pattern and process in
the terrestrial environment. Featuring review questions at the end of
each chapter, suggestions for recommended reading, and a glossary
of ecological terms, Principles of Terrestrial Ecosystem Ecology is
an important text suitable for use in all courses on ecosystem ecology.
Resource managers, land use managers, and researchers will also
welcome its thorough presentation of ecosystem essentials.
2002/472 PP., 199 ILLUS./SOFTCOVER/$52.95
ISBN 0-387-95443-0
ECOREGION-BASED DESIGN
FOR SUSTAINABILITY
ROBERT G. BAILEY, USDA Forest Service, Fort Collins, CO
Illustrations by LEV ROPES
“Bob Bailey [is] the man behind the
ecosystem mapping of the world.”
—LINGUA FRANCA
This richly illustrated volume completes
Robert G. Bailey’s celebrated study of
ecoregions, begun in the landmark Ecosystem
Geography (1996) and further articulated in
Ecoregions (1998). In this third installment,
the author expands his system for defining
large-scale ecological zones to encompass
principles of land management, regional
planning, and design. In an engaging, non-technical discussion, he
shows how larger patterns and processes that characterize a region—its
climate, topography, soils, vegetation, fauna, and human culture—provide
essential keys to the sustainability of ecosystems. Ecoregion-Based
Design for Sustainability will be welcomed by land and resource
managers, landscape architects and urban planners, ecologists, students, and anyone interested in ecology-based design.
2002/240 PP., 100 ILLUS.
AVAILABLE IN SOFTCOVER AND HARDCOVER:
SOFTCOVER/ISBN 0-387-95430-9/$49.95
HARDCOVER/ISBN 0-387-95429-5/$119.00
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2003, VOLUME 6, 8 ISSUES TOTAL
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Promotion S6030
GUIDELINES FOR WRITING PAPER SUMMARY FOR BIO 101
You must turn in a copy of the entire article you are summarizing, either via email (preferably) or as a
printout. Your summary may be emailed or turned in as a hard copy. All emailed materials must be
received at or before the beginning of class on the due date.
Writing the summary:
• The summary should be typed. It must be no longer than one page. There should be no cover
sheet.
• You must summarize each of the sections (Introduction, Methods, Results, Discussion) of the
primary article.
• I do not expect you to understand every detail in the paper, but you must be able to understand
the general principles and research involved. If you don’t understand anything in the paper, then
choose another one to summarize.
• You must use your own words. Do not simply take sentences directly from the paper and put
them in your summary.
• Scientific names must be italicized or underlined. The genus should be capitalized, the species
should not be capitalized. ex: Thalassia testudinum.
• There should be no spelling, grammatical, and typographical errors.
Name: __________________________________________________________________
Grading Criteria for Summary of Scientific Paper
Demonstration of scientific
literacy
Points
Awarded:_____
Demonstration and
documentation of scientific
observations
Points Awarded:_____
Application of the steps in the
scientific method
Points Awarded:_____
Demonstration of analytical
thought through clear conclusions
Points Awarded:_____
Demonstration of written
communication skills; spelling,
grammar, and format
Points
Awarded:_____
Points = 5
Points = 4
Points = 3
Points = 2
Points = 1
The summary
makes it very
clear that the
student
understands the
purpose of the
paper.
There is
evidence that
the student
clearly
understands the
concept of the
paper and the
significance of
the data
provided.
The summary
makes it
sufficiently clear
that the student
understands the
purpose of the
paper.
There is
evidence that
the student
adequately
understands the
concept of the
paper and the
significance of
the data
provided.
The summary
makes it
somewhat clear
that the student
understands the
purpose of the
paper.
There is
evidence that
the student
doesn’t fully
understand
some concepts
of the paper and
the significance
of the data
provided.
The summary
does not make it
clear that the
student
understands the
purpose of the
paper.
There is
evidence that the
student has
minimal
understanding of
the concepts of
the paper and
the significance
of the data
provided.
The summary of
the paper is
vague. There is
evidence that the
student has no
understanding of
the significance
of the information
provided in the
scientific paper.
Scientific facts
and
observations
are present in
the summary,
and are relevant
and correct as
stated.
Scientific facts
and
observations
are present in
the summary,
and mostly
relevant and
correct.
Scientific facts
and
observations are
present in the
summary, but
several are
irrelevant or
incorrectly
stated.
Very few
scientific facts or
observations are
present or
correctly stated
in the summary.
No relevant
scientific facts or
observations are
present or
correctly stated
in the summary.
Steps of the
scientific
method used in
the paper are
defined explicitly
in the summary
using
appropriate
scientific terms.
All issues are
addressed
thoroughly.
Steps of the
scientific
method used in
the paper are
defined
adequately in
the summary.
Some issues
are not
addressed
thoroughly.
Steps of the
scientific
method used in
the paper are
defined
satisfactorily in
the summary.
Several issues
are not
addressed
thoroughly.
Steps of the
scientific method
used in the paper
are not well
defined in the
summary. Most
issues are not
addressed
thoroughly.
Steps of the
scientific method
used in the paper
are not defined in
the summary.
Concepts are
clearly stated
and expressed
thoroughly in
the summary;
analysis is
logical and
complete.
Concepts are
stated and
expressed
adequately in
the summary;
analysis is
mostly logical
and complete.
Concepts are
stated but not
expressed
thoroughly in the
summary;
analysis is
logical but
incomplete.
Concepts are not
stated or are
unclear in the
summary;
analysis is
absent.
Writing is
excellent, word
usage, spelling,
grammar and
punctuation is
excellent.
Formatting is as
instructed.
Writing is
adequate
sufficient use of
wording,
grammar and
punctuation,
very few errors.
Formatting is as
instructed.
Writing is
satisfactory,
average use of
wording,
grammar and
punctuation,
several errors;
Formatting
mostly as
instructed.
Concepts are
minimally stated
and not
expressed
thoroughly in the
summary;
analysis is not
logical and is
incomplete.
Writing is below
average,
insufficient use of
wording,
grammar and
punctuation,
many errors.
Formatting is
mostly not as
instructed.
Writing is poor,
too many
deficiencies in
word use,
grammar,
punctuation and
presentation.
Formatting is not
as instructed.
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