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siderable and increasing stress from human activities (Hutchings, 2000; Jackson et al.,
e
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2001). This unprecedented strain on both the structure and function of marine ecosystems has
v
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led to calls for new management approaches to counter anthropogenic impacts in the coastal ocean
o
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(Botsford et al., 1997; Browman and Stergiou, 2004: Pikitch et al., 2004). Spatial management, includ-
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d t
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the need for models that capture the spatial dynamics of marine populations, especially with respect
g o
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to larval dispersal (Willis et al., 2003; Sale et al., 2005). Theoretical studies suggest that population con-
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structure, genetic diversity, and the resiliency of populations to human exploitation (Hastings and
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Harrison, 1994; Botsford et al., 2001). Modeling efforts have been hindered, however, by the paucity of
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empirical estimates of, and knowledge of the processes controlling, population connectivity in ocean
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ecosystems. While progress has been made with older life stages, the larval-dispersal component of
d r
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connectivity remains unresolved for most marine populations. This lack of knowledge represents a
a
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marine organisms. Furthermore, a lack of spatial context that such information would provide has
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20849-1931,
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14
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The spatial extent of larval dispersal
Finally, estimates of larval dispersal using
interact on different spatial and tempo-
in marine systems has traditionally been
advection/diffusion models with realistic
ral scales to disperse the larvae of marine
inferred from estimates of pelagic dura-
mortality terms and vertical positioning
organisms. Furthermore, a mechanis-
tions of larval dispersive stages, from the
behavior show more restricted move-
tic understanding generates testable
modeled movements of passive particles
ment than would be predicted from one-
hypotheses of larval transport and
by ocean currents, or from analyses of
way oceanic currents acting on passive
dispersal in new environments or loca-
variation in allele frequencies of mito-
particles (e.g., Cowen et al., 2006). Taken
tions. The combination of marker and
chondrial or nuclear genes (Johnson,
together, these studies provide intrigu-
process-oriented approaches promises a
1960; Scheltema, 1988; Planes, 2002).
ing, albeit incomplete, evidence that
truly predictive understanding of larval
Observations of pelagic larval durations
subpopulations of marine organisms
dispersal and connectivity.
(PLDs) of many weeks to over one year
may be more isolated over smaller spatial
The dominant scales of larval disper-
in numerous marine species, coupled
scales than was previously thought. We
sal in coastal species are not known, and
with predicted advection of passive
are, nonetheless, a long way from a com-
perceptions on this issue vary broadly
particles by mean, low-frequency cur-
prehensive understanding of population
within the academic community; opin-
rents, imply that long-distance dispersal
connectivity that would allow for quan-
ions range from broad to restricted
among subpopulations may be perva-
titative predictions of specific natural or
dispersal and from devout to agnostic.
sive. A number of studies documenting
human impacts on marine populations.
The few studies where natal origins have
genetic homogeneity over regional to
Fundamental knowledge of larval dis-
been empirically determined (Jones et
basin-wide spatial scales provides fur-
persal and connectivity can be gained
al., 1999, 2005; Almany et al., 2007), and
ther support for the existence of disper-
from (1) understanding the biological
the case of endemic species on isolated
sal over long distances (e.g., Shulman
and hydrodynamic processes involved
islands where larvae must have origi-
and Birmingham, 1995). More recent
in the transport of larvae and (2) deriv-
nated from local sources (Robertson,
research and careful reconsideration
ing larval origins and dispersal pathways
2001), demonstrate that limited dis-
of the evidence, however, suggests this
using geochemical, genetic, or artificial
persal occurs in marine environments.
perception is likely inaccurate for many
species, particularly over time scales of
ecological relevance.
New hypervariable nuclear DNA
...these papers...set the stage for a groundswell
assays show genetic differentiation
of interdisciplinary scientific and community
among subpopulations of marine fish
and invertebrates that were undetected
interest in marine population connectivity.
by earlier, less-sensitive DNA analyses
(Bentzen et al., 1996; Purcell et al., 2006;
Gerlach et al., 2007). Novel tagging
approaches demonstrate the potential
markers. Natal origins and destination
In contrast, observations that larvae of
for local retention of reef fish larvae
points provide the basic data in connec-
shallow-water species are found in ocean
(Jones et al., 1999, 2005; Almany et al.,
tivity studies (Box 1). However, a pro-
gyre systems, and examples of significant
2007), while constrained nearshore lar-
cess-based understanding of dispersal is
range extensions during narrow event
val distributions of littoral invertebrate
an essential component of population
windows, indicate dispersal on the scale
species (Barnett and Jahn, 1987) suggest
connectivity because it addresses how
of hundreds to thousands of kilometers
localized retention in nearshore waters.
biological and hydrodynamic processes
is also possible (Johnston, 1960; Cowen,
1 Population connectivity refers to the exchange of individuals among geographically separated subpopulations that comprise a metapopulation. Set in the context of benthic-oriented marine
species, population connectivity encompasses the dispersal phase from reproduction to the completion of the settlement process (including habitat choice and metamorphosis).
Oceanography September 2007
15

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1985; Sheltema, 1986; Victor, 1986;
contribution to population replenish-
and diffusive processes that relate to the
Newman and McConnaughey, 1987).
ment and maintenance.
dispersal and recruitment of marine
Identification of relevant temporal
Estimating population connectiv-
organisms is potentially large, sev-
scales is also of critical importance to any
ity in marine ecosystems is inher-
eral general observations may help to
discussion of population connectivity.
ently a coupled bio-physical problem.
define the connectivity problem. First,
For population maintenance, and associ-
Important physical processes include
temporal and spatial correlation scales
ated conservation and resource-manage-
boundary layer structure, particularly
over continental shelves are often quite
ment objectives, the relevant time scale
over the inner shelf, tides, internal tides
short--on the order of days and kilo-
is ecological or demographic, rather than
and bores, fronts and associated jets,
meters. Unfortunately, correlation scales
that relevant to evolutionary processes.
island wakes, and cross-shelf forcing via
near islands, reefs, and within estuaries
Rates of exchange necessary to impact
eddies, meanders, and lateral intrusions.
are not well known. Careful selection
populations on ecological time scales
However, physical processes alone do
of sampling strategies is therefore nec-
are several orders of magnitude higher
not determine the scales of connectivity.
essary to resolve the physical processes
than those required to influence genetic
Time scales of larval development and
described above. Second, the relative
structure. Consequently, both the time
behavioral capabilities, including vertical
contributions of these processes will
over which dispersal is measured and the
migration, also play an important role
likely change from site to site, depend-
amplitude of the relevant recruitment
(Cowen, 2002).
ing on such factors as coastal geometry,
signal must be appropriate for ecological
Although the number of advective
proximity to estuaries, water-column
BOx 1. QuANtitAtiVe MeASureS OF POPulAtiON CONNeCtiVity
A mechanistic understanding of marine population connectivity requires
erature where the basic description of dispersal is a dispersal curve, a one-
resolution of the biological and physical processes involved in larval
dimensional representation of the number of settlers from a given source
dispersal and transport. Larval dispersal refers to the intergenerational
as a function of the distance from that source (see Figure A-1).
spread of larvae away from a source to the destination or settlement site
The dispersal curve becomes a dispersal kernel with an associated
at the end of the larval stage. This usage is widespread in the terrestrial lit-
probability density function, in n dimensions. Formally, the dispersal ker-
nel is the probability of ending up at position x given a starting position y.
One quantitative measure of population connectivity is the source distri-
bution matrix , which gives the proportion of juveniles in population i
ij
Local Retention (Closed)
Broadly Dispersed (Open)
that came from population j. in the absence of any data, let's assume that
larval production in a population is a function of habitat area and that
Population Connectivity
recruitment decays exponentially with distance from a natal population.
in this case,
Figure A-1. One-dimensional, theoreti-
cal dispersal curves depicting dispersal
from a source location ranging from
strong retention to broadly dispersed.

where d is the distance between population i and j, A is the area inhab-
ij
j
ited by population j, and scales the effect of distance on dispersal
(Moilenan and Niemanen, 2002). Although simplistic, the model may
provide an adequate representation of connectivity in metapopulations
dominated by self-recruitment (Jones et al., 2005; Almany et al., 2007).
Distance from Source
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stratification, and seasonal wind forc-
highlights recent advances, as well as
how connectivity influences the dynam-
ing (e.g., Werner et al., 1997; Epifanio
challenges facing the oceanography com-
ics of affected populations. Resolving the
and Garvine, 2001; Sponaugle et al.,
munity, as ocean ecologists seek a mech-
mechanisms controlling larval dispersal
2002; Pineda and Lopez, 2002). Third,
anistic understanding of marine popula-
will involve a coherent understanding
the individual processes contain length
tion connectivity. The major challenges
of the relevant physical processes and
and time scales that vary, and so physi-
in this effort are to provide a quantita-
how organisms mediate the physical
cal transport and dispersal is inherently
tive understanding of the processes and
outcome. Multiple scales will be impor-
a multiscale process. This variability
scales controlling larval dispersal and
tant, and therefore understanding how
presents problems for modeling, as it is
difficult at the present time to resolve
rOBert K. COweN (rcowen@rsmas.miami.edu) is Maytag Professor of Ichthyology and
mesoscale and small-to-intermediate
Chair, Division of Marine Biology and Fisheries, Rosenstiel School of Marine and Atmospheric
scales simultaneously. Finally, there is a
Science, University of Miami, Miami, FL, USA. GleN GAwArKiewiCz is Senior Scientist,
need for a higher degree of precision in
Physical Oceanography Department, Woods Hole Oceanographic Institution, Woods
knowledge of the flow fields in order to
Hole, MA, USA. JeSúS PiNedA is Associate Scientist, Biology Department, Woods Hole
embed behavioral models on particles
Oceanographic Institution, Woods Hole, MA, USA. SiMON r. thOrrOld is Associate
within physical models to test hypoth-
Scientist, Biology Department, Woods Hole Oceanographic Institution, Woods Hole, MA,
eses involving bio-physical interactions.
USA. FrANCiSCO ("CiSCO") e. werNer is George and Alice Welsh Professor, Department
This special issue of Oceanography
of Marine Sciences, University of North Carolina, Chapel Hil , NC, USA.
Physical and coupled bio-physical hydrodynamic models can provide
a more sophisticated parameterization of connectivity models. here,

where p represents the probability that a larva produced in population i
ij
settles in population j (Figure A-2). These probabilities are generated
by coupling output of a hydrodynamic model with lagrangian particle-
tracking protocols that allow for virtual larvae to be assigned variable
pelagic larval durations, vertical migration behaviors, and horizontal
swimming abilities (e.g., Paris et al., in press). By using an individual-based
approach, coupled bio-physical models have flexibility to incorporate
characteristic life-history traits and behavioral capabilities of different
taxa. however, to compare predictions from the various connectivity
Figure A-2. two-dimensional dispersal kernels calculated from a
series of model runs using a coupled biological and physical
models, we need empirical estimates of larval dispersal to evaluate model
model (Cowen et al., 2006; Paris et al., in press). Scale rep-
performance. while new larval mark-recapture approaches are providing
resents probability of successful dispersal from release
information on levels of self-recruitment to local populations, tracking
sites indicated by red dots. Figure provided by
larvae that disperse away from natal locations defines the critical chal-
C. Paris, University of Miami
lenge for field ecologists studying connectivity in marine systems.
Oceanogr
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e te
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17

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OCEANUS
Figure 1. Population connectivity of benthic marine organisms occurs primarily during the pelagic larval phase when individuals either return to their natal loca-
tion to settle, or disperse and settle some distance away from their natal population. while these larval movements are currently shrouded in mystery, new tech-
nologies promise to transform our understanding of population connectivity in ocean ecosystems. For instance, autonomous underwater vehicles (AuVs) could
provide almost continuous real-time data on local hydrography that would then be streamed and assimilated into a coupled bio-physical model to predict the
location of larvae spawned at a particular site. Model predictions could then be relayed to a research vessel conducting adaptive larval sampling using new in
situ imaging systems that would, in turn, provide near-real-time distributions of target larvae. These distributions could then be used to optimize new mission
targets for the AuVs during the following data-collection cycle.
the processes are coupled across scales is
observation, explanation, consequences,
learn into societal gains? Progress has
essential. Identifying patterns will need
and application. These issues can be
been made in all four categories, but in
to involve efforts that focus on a variety
captured, respectively, in the follow-
most cases only at the periphery of the
of species with different life histories
ing general questions: (1) What is the
problem. This may be especially true
across diverse environments. In con-
spatial/temporal distribution of suc-
of the second question, where answers
cert, the problem is multidisciplinary,
cessful settlers originating from source
are likely to be particularly challenging
but one requiring interdisciplinary
populations? (2) What processes influ-
because a variety of physical and biologi-
research effort (Figure 1).
ence the shape of this dispersal kernel?
cal components contribute to the shape
The core challenges or issues rel-
(3) How do connectivity rates influence
of the dispersal kernel. Although these
evant to population connectivity can
population and community dynam-
components can be addressed separately,
be parsed into four specific categories:
ics? (4) How do we translate what we
they will ultimately need to be examined
18
Oceanography Vol. 20, No. 3

---

together due to the role of interactions.
phy of the coastal ocean, with an explicit
and offer broad theoretical contexts for
Ultimately, a process-oriented under-
perspective to physical processes poten-
addressing population ecology issues.
standing is a prerequisite to achieving
tially important to connectivity. Werner,
Jones, Srinivasan, and Almany evalu-
prognostic capability of marine-organ-
Cowen, and Paris examine the state of
ate the significance of connectivity to
ism larval dispersal.
biophysical modeling as it pertains to
the conservation of marine biodiversity.
The series of papers in this volume
connectivity, emphasizing both the capa-
They provide recent evidence that the
demonstrates broad recognition of the
bilities of the models and the assump-
resiliency of marine populations to
relevance of and an active interest in the
tions (i.e., limitations) and pointing to
human exploitation may be linked to
study of population connectivity across
areas of process-oriented research that
species richness, thereby highlighting the
ocean science disciplines. These articles
are required to improve coupled models.
importance of maintaining biodiversity
highlight the importance of spatio-
Hedgecock, Barber, and Edmands dis-
in marine communities. This theme is
temporal scales at a generally finer scale
cuss the potential role and limitations of
further discussed in the final paper by
than previously considered in current
genetic methodologies in assessing popu-
Fogarty and Botsford, who look into the
hydrodynamic models and cross-shelf
lation connectivity. These authors pro-
central role of dispersal and connectiv-
processes. The role of biological fac-
vide a dose of realism regarding the capa-
ity in the dynamics of exploited marine
tors, such as larval behaviors that medi-
bilities of genetic methods for inferring
systems. They discuss the critical impor-
ate the outcome of physical mixing and
connectivity, but also a sense of optimism
tance of understanding dispersal pro-
dispersal, is also evident. Similarly, the
with the incorporation of newer inte-
cesses controlling both larval export and
application of new methodologies (and
grative approaches. Similarly, Thorrold,
movement of later life-history stages in
the need for development of others) sug-
Zacherl, and Levin examine new methods
the specification of effective spatial man-
gests exciting results and the potential
for direct measurements of connectiv-
agement strategies with an emphasis on
for a transformative understanding of
ity in the field using natural and artificial
marine reserves.
the importance of spatial processes in
tags. Their work focuses on geochemi-
In summary, while these papers only
marine systems. As the processes and
cal signatures that exist within calcified
touch on the scope of current work
scale of connectivity are better under-
structures of many marine organisms.
addressing various aspects of popula-
stood, the applications of these findings
The last three papers explore the
tion connectivity in marine populations,
are also being dissected to enhance man-
various implications and applications of
they set the stage for a groundswell of
agement and conservation measures.
Each paper in this issue addresses
the current state of knowledge, new
and novel methods for studying
Our hope is that through this combined effort,
connectivity-related processes, and a


oceanographers may be able to establish
call for future work to bring the whole
problem into focus. The first paper, by
a simplified yet useful set of guidelines...
Pineda, Hare, and Sponaugle, discusses
larval transport and larval dispersal and
how they relate to population connectiv-
ity. The authors consider the concept of
connectivity in marine systems. Gaines,
interdisciplinary scientific and commu-
population connectivity, with an empha-
Gaylord, Gerber, Hastings, and Kinlan
nity interest in marine population con-
sis on understanding the role of plank-
discuss the observational and theoreti-
nectivity. Our hope is that through this
tonic processes on the success of the
cal advances and challenges in under-
combined effort, oceanographers may be
settlers. Gawarkiewicz, Monismith, and
standing the population consequences
able to establish a simplified yet useful
Largier explore the physical oceanogra-
of larval dispersal and connectivity,
set of guidelines (e.g., certain biologi-
Oceanography September 2007
19

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cal processes, such as vertical behavior
opment of this special issue. We also
tem off Southern California. Continental Shelf
Research 7:1­25.
by larvae, may mediate or simplify the
acknowledge the help we received in
Bentzen, P., C.T. Taggart, D.E. Ruzzante, and D. Cook.
dispersive complexity of the physical
all facets of producing this issue from
1996. Microsatellite polymorphism and the popu-
environment). Until we do so, we may
the editor of Oceanography, Ellen
lation structure of Atlantic cod (Gadus morhua)
in the northwest Atlantic. Canadian Journal of
be relegated to resolving connectiv-
Kappel. Over the years, we have each
Fisheries and Aquatic Sciences 53:2,706­2,721.
ity individually for every species and
received support from a variety of agen-
Botsford, L.W., J.C. Castilla, and C.H. Peterson. 1997.
The management of fisheries and marine ecosys-
system of interest.
cies in support of research relevant to
tems. Science 277:509­515.
Population Connectivity; in addition to
Botsford, L.W., A. Hastings, and S.D. Gaines. 2001.
ACKNOwledGeMeNtS
NSF, we acknowledge funding from the
Dependence of sustainability on the configuration
of marine reserves and larval dispersal distance.
The authors would like to acknowl-
World Bank/GEF Coral Reef Targeted
Ecology Letters 4:144­150.
edge many of our colleagues who have
Research Program, University of Miami's
Browman, H.I., and K.I. Stergiou. 2004. Perspectives
on ecosystem-based management approaches to
shared their ideas on population con-
Maytag Chair in Ichthyology, the Oak
the management of marine Resources. Marine
nectivity in a variety of forums over
Foundation, and the Woods Hole
Ecology Progress Series 274:269­270.
Cowen, R.K. 1985. Large scale pattern of recruit-
the last decade. We appreciate the sup-
Oceanographic Institution.
ment by the labrid, Semicossyphus pulcher: Causes
port we have received from the National
and implications. Journal of Marine Research
Science Foundation (NSF). In particu-
reFereNCeS
43:719­743.
Cowen, R.K. 2002. Larval dispersal and retention and
lar, we acknowledge support from Phil
Agardy, T.S. 1997. Marine Protected Areas and Ocean
consequences for population connectivity. Pp. 149­
Conservation, R.G. Landes Co., Austin, TX.
Taylor (Biological Oceanography) and
170 in Ecology of Coral Reef Fishes: Recent Advances.
Almany, G.R., M.L. Berumen, S.R. Thorrold, S. Planes,
P.F. Sale, ed., Academic Press, San Diego, CA.
Eric Itswiere (Physical Oceanography)
and G.P. Jones. 2007. Local replenishment of coral
Cowen, R.K., C.B. Paris, and A. Srinivasan. 2006.
for funding the NSF Workshop on
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