Articles

The Identification,
Conservation, and
Management of Estuarine
and Marine Nurseries for

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Fish and Invertebrates
MICHAEL W. BECK, KENNETH L. HECK, JR., KENNETH W. ABLE, DANIEL L. CHILDERS, DAVID B. EGGLESTON,
BRONWYN M. GILLANDERS, BENJAMIN HALPERN, CYNTHIA G. HAYS, KAHO HOSHINO, THOMAS J. MINELLO,
ROBERT J. ORTH, PETER F. SHERIDAN, AND MICHAEL P. WEINSTEIN

N earshore estuarine and marine ecosystems—e.g.,

seagrass meadows, marshes, and mangrove forests—
serve many important functions in coastal waters. Most no-
A BETTER UNDERSTANDING OF THE
tably, they have extremely high primary and secondary pro-
HABITATS THAT SERVE AS NURSERIES FOR
ductivity and support a great abundance and diversity of MARINE SPECIES AND THE FACTORS THAT
fish and invertebrates. Because of their effects on the diver-
sity and productivity of macrofauna, these estuarine and CREATE SITE-SPECIFIC VARIABILITY IN
marine ecosystems are often referred to as nurseries in nu-
merous papers, textbooks, and government-sponsored re- NURSERY QUALITY WILL IMPROVE
ports (Boesch and Turner 1984, NRC 1995, Butler and Jer-
nakoff 1999). Indeed, the role of these nearshore ecosystems
CONSERVATION AND MANAGEMENT OF
as nurseries is an established ecological concept accepted by THESE AREAS
scientists, conservation groups, managers, and the public

Michael Beck is director of the Coastal Waters Program at The Nature Conservancy and a research associate in the Institute of Marine Sciences,
Center for Ocean Health, 100 Shaffer Road, University of California, Santa Cruz, CA 95060. Kenneth Heck is a professor at the Dauphin Island
Sea Lab and University of South Alabama, Dauphin Island, AL 36528. Kenneth Able is director of the Rutgers University Marine Field Station, Tuck-
erton, NJ 08087, and professor in the Institute of Marine and Coastal Sciences, Rutgers University. Daniel Childers is associate professor in the
Department of Biological Sciences, Southeast Environmental Research Center, Florida International University, Miami, FL 33199. David Eggleston
is associate professor in the Department of Marine, Earth and Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695-8208.
Bronwyn Gillanders is a postdoctoral research fellow in the Department of Environmental Biology at the University of Adelaide, SA, 5005, Australia.
Benjamin Halpern and Kaho Hoshino are graduate students at the University of California, Santa Barbara, CA 93106; Halpern is in the Depart-
ment of Ecology, Evolution, and Marine Biology, and Hoshino is in the Bren School for the Environment. Cynthia Hays is a graduate student in the
Department of Biology, University of California, Santa Cruz, CA 95064. Thomas Minello is chief of the Fishery Ecology Branch, and Peter Sheridan
is a research ecologist in the National Marine Fisheries Service, at Southeast Fisheries Science Center Laboratory, National Marine Fisheries Ser-
vice, 4700 Avenue U, Galveston, TX 77551. Robert Orth is a professor in the School of Marine Science, Virginia Institute of Marine Science, Col-
lege of William and Mary, Gloucester Point, VA 23062. Michael Weinstein is director of the New Jersey Marine Sciences Consortium, Fort Hancock,
NJ 07732. © 2001 American Institute of Biological Sciences.

August 2001 / Vol. 51 No. 8 • BioScience 633

Articles

and cited as justification for the protection and conservation Childers et al. 2000). This transfer of productivity from
of these areas. Nonetheless, the nursery-role concept has coastal ecosystems to food webs is undoubtedly important.
rarely been stated clearly, even in papers that purport to test Nonetheless, there is a separation in the conceptual under-
it. This ambiguity hinders the effectiveness of the nursery-role pinnings and testing of hypotheses about the effects of ecosys-
concept as a tool for conservation and management. We seek tems on the productivity of individual species versus their ef-
to redress that ambiguity by briefly tracing the history of the fects on the productivity of estuaries and coastal oceans in
concept, developing a clear hypothesis with testable predic- general. An analysis of these effects is beyond the scope of this
tions, and discussing how this work can focus efforts in re- paper, but they will be addressed in a future work.
search, conservation, restoration, and management. Most studies of the nursery-role concept have examined the
effects of seagrass meadows or wetlands on either the density,
History of the nursery-role concept survival, or growth of juveniles on the species’ movement to
The nursery-role concept was first applied nearly a century adult habitats (Figure 1; Heck et al. 1997, Butler and Jer-

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ago to motile invertebrates and fishes with complex life cy- nakoff 1999, Minello 1999). Some studies make direct com-
cles, in which larvae are transported to estuaries, metamor- parisons of these parameters among the habitats used by a
phose, grow to subadult stages, and then move to adult habi- species (Weinstein and Brooks 1983, Sheridan 1992, Jenkins
tats offshore. Gunter (1967) traces this idea to work on blue and Wheatley 1998), but such comparisons are often limited
crabs on the Atlantic coast of the United States (Hay 1905), to vegetated versus unvegetated habitats (Edgar and Shaw
penaeid shrimp on the Gulf of Mexico coast, and finfish on 1995, Gray et al. 1996). Generally, an area has been called a
both of these coasts (Hildebrand and Schroeder 1928). The nursery if a juvenile fish or invertebrate species occurs at
concept became so pervasive that it has been termed a “law” higher densities, avoids predation more successfully, or grows
(Gunter 1967). For example, Deegan (1993, p. 74) states that faster there than in a different habitat.
“estuarine fish faunas around the world are dominated in Of all the studies on the nursery-role concept, most have
numbers and abundance by species which move into the es- focused on the effects of seagrass meadows or wetlands on
tuary as larvae, accumulate biomass, and then move off- an animal’s density. The evidence usually indicates that the
shore.” density of fish and invertebrates is higher in vegetated than
In early papers the estuary as a whole was considered to be in unvegetated habitats (for reviews see Orth et al. 1984,
the nursery. In subsequent works, however, the focus shifted Heck et al. 1997, Able 1999, Minello 1999). Direct compar-
to specific areas within estuaries as nurseries, especially wet- isons of an animal’s abundance between mangrove forests and
lands (herein marshes and mangrove forests) and seagrass other habitats are rare (Sheridan 1992). There are also dif-
meadows, because evidence suggested that they supported ficulties with these comparisons, because different sampling
much greater densities of organisms than adjacent unvege- methods usually are used to estimate densities inside and out-
tated (i.e., without macrophytes) substrates (Williams 1955, side of mangrove forests and frequently samples are only col-
Hutchings and Recher 1974, Turner 1977, Orth et al. 1984, lected in areas adjacent to mangrove forests rather than di-
Minello 1999). We concentrate on seagrass meadows and rectly within the flooded forest.
wetlands because most research to date has addressed their The few studies that have focused on differences in juve-
potential to serve as nurseries. Examples are drawn from nile survival among wetlands, seagrass meadows, and other
other ecosystems when possible and we note that the poten- areas indicate that survival of a species is generally greater in
tial nursery value of some of them, for example oyster reefs, vegetated than in unvegetated habitats (Orth et al. 1984,
has not received due recognition. Throughout the paper, the Heck and Crowder 1991, Able 1999). Even fewer studies
term ecosystem is used to identify characteristic assemblages have focused on the effects of wetlands and seagrass mead-
of plants and animals (e.g., marshes or oyster reefs). The ows on the growth of fish and invertebrates (Heck et al.
term habitat refers to the area used by a species, with modi- 1997, Phelan et al. 2000). In seagrass meadows, evidence re-
fiers added to identify the particular habitats used by an an- garding growth is, surprisingly, equivocal. Only about half of
imal. For example, the blue crab, Callinectes sapidus, has a sea- the studies report that the growth rate of individuals is
grass habitat and a marsh habitat, which refer to particular higher in seagrass habitats than in adjacent habitats (Heck
portions of seagrass and marsh ecosystems, respectively, used et al. 1997).
by the crab. Finally, only a handful of studies have attempted to deter-
We also focus on the direct effects of ecosystems on the pro- mine whether the juveniles of a species move successfully from
ductivity of individual species as opposed to their contribu- putative nursery habitats to adult habitats (Costello and Allen
tions to the productivity of coastal oceans. Seagrass meadows 1964, Deegan 1993, Gillanders and Kingsford 1996, Gillan-
and wetlands have been identified as nurseries in part because ders 1997, Fry et al. 1999). The evidence that supports suc-
they export vast quantities of carbon, nitrogen, and phos- cessful movement of seagrass- or wetland-associated juveniles
phorus to coastal food webs. This export may occur through to adult habitats is largely indirect (Eggleston 1995), both be-
the direct transfer of animal biomass via movement of indi- cause movement data are difficult to obtain and because
viduals, predation, or outwelling of dissolved and particulate there has been a dearth of communication between benthic
organic matter (Teal 1962, Nixon 1980, Deegan 1993, Lee 1995, ecologists and fisheries biologists.

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Figure 1. Examples of field experiments and observations used to assess whether some habitats serve as nurseries. (a) A drop
trap used to compare density between marsh and nearby unvegetated habitats. (b) A tethered shrimp used to assess
differences in survival between sand, seagrass, and marsh habitats. (c) Cages used to examine shrimp growth between marsh
and unvegetated habitats. (d) A juvenile summer flounder, Paralichthys dentatus, being injected subcutaneously with a
nontoxic acrylic paint marker to examine movement patterns.

There is growing recognition that there are exceptions to ports or refutes the nursery-role concept is exacerbated by the
the nursery-role concept. For example, few commercially fact that the nursery-role concept does not have a clearly de-
important species of fish and invertebrates appear to rely fined hypothesis, and therefore it has been difficult to test di-
exclusively on seagrass meadows in coastal waters of Massa- rectly (Edgar and Shaw 1995, Gillanders 1997).
chusetts (Heck et al. 1995) or New Jersey (Able and Fahay
1998). Instead, most of these species use seagrass meadows op- A nursery-role hypothesis
portunistically but can survive well in unvegetated areas. The underlying premise of most studies that examine
Edgar and Shaw (1995) reported that seagrass beds in south- nursery-role concepts is that some nearshore, juvenile habi-
ern Australia were not always better nurseries than nearby un- tats contribute disproportionally to the production of indi-
vegetated substrates. A study on the labrid Australian blue viduals that recruit to adult populations. From this premise,
groper, Achoerodus viridis, indicated that recruits to the off- we have developed a hypothesis from which clear and testable
shore adult population came primarily from young that set- predictions can be made: A habitat is a nursery for juveniles
tled in offshore rocky reefs, not from the abundant young in of a particular species if its contribution per unit area to the
inshore seagrass beds (Gillanders and Kingsford 1996). A production of individuals that recruit to adult populations is
recent planning document produced for the Australian Fish- greater, on average, than production from other habitats in
eries Research Development Corporation concluded that which juveniles occur.
there was very little strong evidence that Australian seagrass The ecological processes operating in nursery habitats, as
provided critical nursery habitat for the majority of Aus- compared with other habitats, must support greater contri-
tralian finfish species (Butler and Jernakoff 1999). butions to adult recruitment from any combination of four
That the evidence about the role of certain ecosystems as factors: (1) density, (2) growth, (3) survival of juveniles, and
nurseries is sometimes contradictory is not surprising— (4) movement to adult habitats (Figure 2). A general null hy-
there are exceptions to any broad ecological concept. How- pothesis is that there is no difference in the nursery value (i.e.,
ever, much of the disagreement about evidence that sup- production of individuals that recruit to adult populations per

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Articles

Figure 3. Relationship between juvenile, nursery, and

adult habitats. The square represents all habitats. The
ovals represent the portions of habitats used during
juvenile and adult stages. Nursery habitats are a subset

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of juvenile habitats. (a) Classic concept of species that
have nursery habitats. (b) General concept of species that
have nursery habitats. There can be overlap in the
habitats that juveniles and adults use, but there must be
some movement to nonjuvenile habitats for a species to
be considered to have a nursery habitat. This movement
is often associated with reproduction or an ontogenetic
habitat shift.
mouths of estuaries) to nonjuvenile habitats to release eggs
(Orth and van Montfrans 1990). Some species do not move
Figure 2. A representation of the factors operating in directly from juvenile to adult habitats but move gradually be-
juvenile and nursery habitats. The thickness of the arrows tween them (e.g., spiny lobsters), and they also are considered
indicates the relative contribution from each factor to the to have nursery habitats.
recruitment of adults. A nursery habitat (dashed oval) We suggest that species must have at least some disjunction
supports a greater than average combination of increased between juvenile and adult habitats to be considered to have
density, survival, and growth of juveniles and movement nursery habitats (Figure 3b), and in most of these species,
to adult habitats. (a) All four factors are greater in the movement to nonjuvenile habitat is associated with repro-
nursery versus other juvenile habitats. (b) Only one of the duction. There are many other life history strategies, of
four factors, in this case movement, is greater in the course—this hypothesis does not imply that seagrass mead-
nursery versus other juvenile habitats. ows, for example, do not have important effects on species that
unit area of juvenile habitat) of different juvenile habitats for spend their entire life there. These other life history strategies,
a given species. however, do not fit the nursery-role hypothesis. Based on
our definition, taxa that do not have nurseries per se include,
Considerations for tests of the for example, bay scallops (Argopecten irradians), killifish
nursery-role hypothesis (Fundulus), bay anchovy (Anchoa mitchilli), and amphipods.
There are a number of key considerations on the species, habi- Examples of taxa that do have nurseries are clawed lobster
tats, and variables that should be accounted for when testing (Homarus americanus), eels (Anguillav), red drum (Sciaenops
the nursery-role hypothesis. These considerations have fre- ocellatus), gag grouper, blue groper, pink snapper (Pagrus
quently been overlooked in the past. auratus), luderick (Girella tricuspidata), tarwhine (Rhab-
dosargus sarba), blue crabs, brown shrimp (Farfantepenaeus
The nursery-role hypothesis focuses on a particu- aztecus), flounder (Paralichthys spp.), pinfish (Lagodon rhom-
lar set of life history strategies—that is, on those boides), striped mullet (Mugil cephalus), and gray snapper (Lut-
strategies where there is a separation between juvenile and janus griseus).
adult habitats (Figure 3). The original literature on nurs-
eries focused on an idealized or classic life history strategy: Ju- The nursery role of habitats must be compared
veniles grew up in nearshore or estuarine habitats and then on a unit-area basis. Even if a habitat is small in area, it
undertook rapid, directional movement to completely different is an important nursery habitat if it produces relatively more
offshore adult habitats (Figure 3a). The gag grouper (Myc- adult recruits per unit of area than other juvenile habitats used
teroperca microlepis), for example, fit this classic life history by a species. This distinction is important in conservation and
strategy (Koenig and Coleman 1998). However, many other management, where priorities must be set for limited fund-
species with substantial overlap in juvenile and adult habitats ing and effort. It is more important to conserve, abate the loss,
have historically been thought to use nurseries. In blue crabs, restore, or otherwise manage habitats that contribute dis-
for example, juveniles and adults often occupy the same habi- proportionately to the production of adults. This need is
tats, but females make a directed movement (usually to the even more pressing if these habitats are relatively uncommon.

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It is possible that common habitats may make important

contributions to the recruitment of adults even if the density
of individuals per area is low, simply because the habitats are
widespread. We predict, however, that there will be few cases
where habitats that have lower densities and often lower sur-
vival and growth rates of individuals will make significant con-
tributions to adult recruitment simply because they are wide-
spread. And if these habitats do make significant contributions
solely because of their large areal coverage, they will be im-
portant juvenile habitats, but not nurseries per se.

A definitive test of the nursery-role hypothesis re-

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quires a comparison among all habitats that ju-
veniles use (Figure 4). Comparisons among putative
nursery habitats have usually involved only vegetated and
unvegetated habitats, even though individual species may
use many different habitats (Minello 1999). Thus, seagrasses
or wetlands may seem less important as nurseries in regions
where alternative habitats are used successfully. For example,
in bays in southern Australia and in the northeastern United
States, a species may be found in many habitats (e.g., cobble, Figure 4. A hypothetical comparison of the nursery value
rocky reef, oyster reef, kelp, sandy or muddy bottom) in ad- of several different habitats. The dashed line represents
dition to its marsh and seagrass habitats (Ward et al. 1999). the average percentage productivity of adults per unit
To determine which, if any, habitats serve as nurseries, all of area from all the juvenile habitats. In this example
a species’ juvenile habitats should be surveyed. seagrass meadows, marshes, and oyster reefs are nursery
habitats.
Nursery habitats are a subset of juvenile habitats.
Any habitat that makes a greater than average contribution The total biomass of individuals recruiting to
to the recruitment of adults should be considered a nursery adult populations is the best single measure of the
habitat. Thus, some portions of juvenile habitats will be nurs- contribution from juvenile habitats. The nursery
eries, but not all juvenile habitats can be nurseries (Figure 3). habitats for a species are those that are the most likely to
Previously, there has been little discussion of the quantitative contribute to future populations. This contribution should be
contribution that a habitat must make before it is considered a function of both the size and number of individuals that re-
a nursery. In most tests, however, a habitat was considered a cruit to adult populations, because these variables affect sur-
nursery if some parameter (usually density) was statistically vival, growth, and reproductive success in the adult habitats.
significantly greater in that habitat than in another. This us- Total biomass (i.e., production) of individuals recruiting to
age implies that any habitat with a greater than average con- adult populations should be the best integrative measure of
tribution to adult recruitment should be considered a nurs- this potential contribution from juvenile habitats to future gen-
ery. Juvenile habitats that are found not to be nurseries can erations.
and often do contribute individuals to adult populations,
but they make a less than average contribution when compared Examinations of the density of juveniles among
with other habitats (Figure 4). If many habitats are examined, habitats do not provide a conclusive test of the
it should be possible to identify and focus on those that make nursery-role hypothesis. In the overwhelming majority
the greatest contribution to adult recruitment, that is, the best of studies, a habitat is suggested to be a nursery largely because
nursery habitats. it supports high densities of juveniles relative to another
habitat. It is assumed that higher juvenile densities will lead
The movement of individuals from juvenile to to a greater recruitment to adult populations. Although a
adult habitats must be measured. There are very few habitat may support high densities of juveniles, if these indi-
studies on movement patterns, and this is a vital missing viduals never reach adult populations, then that habitat does
link in our understanding of nurseries. Movement of indi- not function as a productive nursery. In most studies the
viduals is one of the most difficult variables to measure in ecol- unstated premise has been that, all else being equal, habitats
ogy. Fortunately, vast improvements in technology—archival with higher densities of juveniles are likely to make a greater
data loggers, stable isotopes, genetic markers, and otolith mi- contribution to the production of adults than habitats with
crochemistry—allow researchers to track and infer move- lower densities of juveniles. This correlation, which is rarely
ments (Gillanders and Kingsford 1996, Thorrold et al. 1998, tested, may hold in many cases, but there are likely to be im-
Fry et al. 1999). portant exceptions. For example, some sites may be well

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Articles

placed to receive larval influx, but these could be sites where as nurseries; greater attention needs to be paid to their in-
juveniles grow slowly or where movement to adult habitats is teraction with postsettlement processes.
risky or difficult (e.g., there are no adult habitats nearby or Landscape-level factors also can affect the nursery value of
there is particularly intense predation; Lipcius et al. 1997, sites within habitats (Table 1). For example, the relative location
McBride and Able 1998). Density is only one of four factors of seagrass beds in an estuary can affect the density of fish
that must be considered to determine whether a habitat species; some seagrass beds near the site where larvae enter
serves as a nursery. estuaries have higher densities of fish than similar beds far-
It also is not sufficient to measure how long individuals ther up the estuary (Bell et al. 1988). Lipcius and colleagues
spend in nursery habitats to determine whether that habitat (1997) suggested that proximity—i.e., relative location of
is a nursery. That is, the duration of occupancy is important nursery and adult habitats in the Exuma Sound, Bahamas
only inasmuch as it contributes to a greater combination of seascape—affects the abundance of adult lobsters by affect-
survival and growth of the individuals that leave the nursery ing the success of movement between habitats. Relative lo-

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habitat. cation, with respect to large water movements such as up-
welling or retention zones, has also been shown to strongly
Factors that contribute to site-specific influence larval delivery (Roy 1998), thus playing a crucial role
variation in nursery value in setting initial juvenile densities within a habitat. Irlandi and
The nursery value of seagrass meadows, wetlands, and other Crawford (1997) concluded that for pinfish the nursery value
ecosystems may vary geographically. For example, recent of salt marshes was affected by their location relative to sea-
analyses suggest that seagrass meadows in the tropical grass beds: Both the density and growth of pinfish were
Caribbean are more important as nurseries than they are in higher in marshes adjacent to seagrass beds than in marshes
the Indo-Pacific region (Williams 1991); other analyses have adjacent to unvegetated bottom. Several good landscape-
found seagrass meadows more important as nurseries in the scale studies document phenomena that are likely to create
United States than in Australia (Edgar and Shaw 1995, But- variation in the value of nursery habitats, even though they
ler and Jernakoff 1999, Ward et al. 1999). Within the United do not specifically address the nursery-role hypothesis. For ex-
States, seagrass meadows in warm temperate regions may ample, Irlandi (1994) found that rates of predation on clams
serve as better nurseries than those in cool temperate re- were higher in more fragmented seagrass beds. Micheli and
gions (Orth and van Montfrans 1990, but see Grant and Peterson (1999) found that densities of macroinvertebrates on
Brown 1998). Marshes in the Gulf of Mexico are suggested to oyster reefs were lower where the reefs were next to salt
be more important as nurseries than marshes in the US South marshes, which harbored blue crab predators. The importance
Atlantic (Minello 1999). of these factors (Table 1) needs to be better examined, because
This potential geographic variation is a source of con- much of the apparent discrepancy in nursery roles in differ-
tention about the importance of nurseries in general. Much ent regions (across latitudinal gradients or between continents)
of the apparent discrepancy in the importance of nurseries in very likely can be attributed to one or several of these factors
different regions could be understood, however, by examin- operating locally (e.g., within estuaries).
ing factors that contribute to local variation (e.g., within
estuaries) in nursery value. For example, even within an Implications for research, conservation,
estuary there is variation in the nursery value of different sea- restoration, and management of
grass meadows for a species. Factors that can create this site- nurseries
specific variation in the nursery value of habitats can be Throughout the world, the degradation of coastal ecosys-
grouped into three broad categories: biotic, abiotic, and land- tems continues at an alarming rate (Hinrichsen 1998). Estu-
scape (Table 1). aries may be some of the most degraded environments on
Many biotic and abiotic factors can influence the nursery
value of habitats for a species (Table 1). For example, Heck and Table 1: Factors that create site-specific variation in
Crowder (1991) found that predation on target species in sea- nursery value
grass beds was lower in more structurally complex beds,
which suggests that more complex beds may serve as better Biotic Abiotic Landscape
nurseries for many species because they increase survivorship.
Salinity also appears to have important effects on site-specific Larval supply Water depth Spatial pattern
Structural complexity Physico-chemical (e.g.,size,
variation in the nursery value of habitats. For example, the den- Predation (dissolved O2, shape,
sities of many species within marshes are highly dependent Competition salinity) fragmentation,
on salinity (Minello 1999). Larval supply and presettlement Food availability Disturbance regime connectivity)
Tidal regime Relative location
processes also can affect the initial density and condition (e.g., to larval
(e.g., size) of juveniles within a habitat (Grimes and Kings- supply, other
ford 1996, Roy 1998). In general, presettlement processes are juvenile habi-
tats, adult habi-
rarely considered when evaluating how well habitats function tats)

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earth, because they have been focal points for human colo- periments are preferred also because they often lead to use-
nization for centuries (Edgar et al. 2000). Interest in conserving ful insights about factor interaction.
and managing coastal waters is intense and widespread, but Second, researchers must consider multiple habitats. Al-
funds are limited and must be targeted judiciously. Devel- though most species are found in more than one or two
opment of a better nursery-role hypothesis may help re- habitats, surprisingly few studies make comparisons among
searchers identify the habitats and, even more important, more than two potential nursery habitats.
the sites within habitats that serve as nurseries for a species, Third, we must attempt to better quantify the movement
thus focusing efforts in research, conservation, restoration, and of individuals between juvenile and adult habitats with all
management. However, it is not useful to wait for irrefutable available tools. Refinements in tagging and chemistry will
evidence of a given area’s function as a nursery before action help substantially to identify the sources of individuals that
is taken to conserve, manage, or restore it. Rather, it is neces- recruit to adult habitats, yet these techniques can be labor in-
tensive and expensive; moreover, they involve more labora-

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sary to err on the side of caution and to act on current knowl-
edge of the potential for a given area to serve as a nursery for tory than field work, which would require a major shift in
some species. many research programs. Nonetheless, it should be possible
Seagrasses and wetlands have been the focus of most work to design simple but elegant field studies to examine the
on nurseries, and in many cases this emphasis is justified. How- movement of juveniles. It is surprising, for example, that so
ever, improved tests of predictions from the nursery-role hy- few studies examine season- and size-specific movements of
pothesis may show that previously ignored areas also serve as juveniles out of the mouths of estuaries towards adult habi-
nurseries and therefore should be better conserved and man- tats (Deegan 1993).
aged (Gray et al. 1996). The question this article addresses is Fourth, although we have focused on direct methods of
not “Are wetlands and seagrasses important?” There is un- study in this article, correlative and case study analyses can yield
deniable evidence of their importance, aside from their po- many useful insights. For example, Butler and Jernakoff
tential as nurseries, at many sites. They provide many ecosys- (1999) reviewed many studies that looked for correlations be-
tween inshore habitat loss and offshore fisheries production.
tem services and serve many important functions (Costanza
These correlative analyses cannot provide strong inference for
et al. 1997), stabilizing shorelines, reducing wave impacts,
the existence of nursery habitats, but they do provide relevant
removing suspended solids, recycling nutrients, and adding
observations on potential nurseries at scales that are ecolog-
oxygen to surrounding waters (Short and Wyllie-Echeverria
ically and economically important.
1996, Costanza et al. 1997, Gosselink et al. 1999). Seagrasses
Better and more consistent tests of the nursery-role hy-
and wetlands are highly productive, and this production en-
pothesis will identify nursery habitats. More important, they
ters coastal food webs through many different pathways, not
will reveal which factors create site-specific variation within
just as fish moving to adult habitats.
habitats in the production of juveniles that recruit to adult
The development of the nursery-role concept is similar in
populations. These tests should also provide a better indica-
some respects to the development of the keystone species tion of the species that depend on particular nursery habitats.
concept. There are few rigorous tests of predictions developed Conservation and management organizations now com-
from the keystone species concept, and it is difficult to con- monly consider all seagrasses and wetlands as nurseries. These
duct all the experiments that would be necessary to show un- broad declarations may be useful for generating public interest,
equivocally that a keystone species exists (Power et al. 1996). but they hinder the actual work that needs to be accom-
Nonetheless, it would be useful to know what a definitive test plished by these groups because the statements lack focus. A
would encompass, so that researchers could arrive at the best clearer understanding of the habitats that serve as nurseries
approximation of it. Although there is no unequivocal test of for species, and the factors that make some sites more valu-
the keystone species concept, sufficient evidence exists to in- able as nurseries, will allow more efficient use of limited
dicate that some species are likely to be keystone species money, time, and effort in conservation and management. Not
(Estes and Duggins 1995) and others are not (Elner and all oyster reefs, cobble, or wetlands are created equal. If it were
Vadas 1990). The situation is much the same for nursery known, for example, that for some species the best seagrass,
habitats. For example, substantial evidence supports the con- marsh, or mangrove nurseries were large areas near sources
tention that some seagrasses and wetlands are likely to serve of larval influx and in close proximity to adult habitats, then
as nurseries (Heck et al. 1997, Butler and Jernakoff 1999, efforts in habitat conservation and management aimed at pre-
Minello 1999) even if there is no definitive test. serving or restoring nurseries could be more judiciously in-
Many practical considerations can help in the testing of pre- vested in those types of sites.
dictions from the nursery-role hypothesis. First, more than one Some of this information is or should be available, but it
factor must be considered. Ideally, all four factors—density, has not been applied specifically to the identification of the
growth, survival, and movement—would be examined in a habitats and the sites within habitats that serve as nurseries.
study of putative nursery habitats, but doing so may be dif- A better understanding and testing of predictions of the
ficult. Nonetheless, researchers cannot continue to be satis- nursery-role hypothesis should enable scientists and funding
fied with single-factor studies in this field. Multifactor ex- agencies to fill the gaps in our knowledge, help nongovern-

August 2001 / Vol. 51 No. 8 • BioScience 639

Articles
Gillanders BM, Kingsford MJ. 1996. Elements in otoliths may elucidate the
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to protect the diversity of species and natural resources, and lations of a temperate reef fish. Marine Ecology Progress Series 141:
allow state and federal agencies and fishery management 13–20.
councils to make better regulatory decisions for fisheries Gosselink JG, Coleman JM, Stewart RE Jr. 1999. Coastal Louisiana. Pages
management, habitat conservation, habitat restoration, and 385–436 in Mac MJ, Opler PA, Puckett Haecker CE, Doran PD, eds. Sta-
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Grant SM, Brown JA. 1998. Nearshore settlement and localized populations
Acknowledgments of age-0 Atlantic cod (Gadus morhus) in shallow coastal waters of New-
This work was conducted as part of the Nursery Roles Work- foundland. Canadian Journal of Fisheries and Aquatic Sciences 55:
ing Group, supported by the National Center for Ecological 1317–1327.
Gray CA, McElligott DJ, Chick RC. 1996. Intra- and inter-estuary differences
Analysis and Synthesis, a center funded by the National Sci-
in assemblages of fishes associated with shallow seagrass and bare sand.
ence Foundation (grant no. DEB-0072909), University of

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