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1/18Oceanography V . 20, N . 322

M a r i N e P o P u l at i o N C o N N e C t i Vi t y

Oceanography V . 20, N . 322

M a N y M a r i N e s P e C i e s have small, pelagic early li e stages. For those spe-cies, knowledge o population connectivity requires understanding the origin andtrajectories o dispersing eggs and larvae among subpopulations. Researchers haveused various terms to describe the movement o eggs and larvae in the marine envi-ronment, including larval dispersal, dispersion, dri t, export, retention, and larvaltransport. Though these terms are intuitive and relevant or understanding thespatial dynamics o populations, some may be nonoperational (i.e., not measur-able), and the variety o descriptors and approaches used makes studies di cult to

compare. Furthermore, the assumptions that underlie some o these concepts arerarely identi ed and tested. Here, we describe two phenomenologi-

cally relevant concepts, larval transport and larval dispersal.These concepts have corresponding operational de nitions,are relevant to understanding population connectivity,and have a long history in the literature, although they are

sometimes con used and used interchangeably. A ter de n-ing and discussing larval transport and dispersal, we consider

the relative importance o planktonic processes to the overallunderstanding and measurement o popula-

tion connectivity. The ideas considered in thiscontribution are applicable to most benthic

and pelagic species that undergo trans orma-tions among li e stages. In this review, however,

we ocus on coastal and nearshore benthicinvertebrates and shes.

l v

t n p nd D pn C oc nC n q nc

P p n C nn cB y J e s s P i N e D a , J o N a t h a N a . h a r e , a N D s u s P o N a u g l e

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2/18Oceanography s p mb 2007 2

l v n pis de ned as the hori-zontal translocation o a larva betweenpoints x1,y 1 and x2,y 2, where x and y arehorizontal axes, say, perpendicular andparallel to the coastline. In larval trans-port, only the spatial dimensions mat-ter. Although this de nition ignoresthe vertical axis (z) or simplicity, thisdimension is critical or larval transportbecause larvae can modi y their hori-zontal distribution by swimming verti-cally, thereby encountering di erentcurrents (Nelson, 1912; Crisp, 1976). Totrans er rom point x1,y 1 to point x2,y 2, alarva can swim horizontally and may betransported by di usive and advectiveprocesses (Scheltema, 1986). De ned asthe translocation o a larva between twopoints, larval transport appears decep-tively simple. However, the wide rangeo larval behaviors and physical mecha-nisms, together with their variability atmultiple scales, makes larval transportexceedingly di cult to measure. Thetemporal and spatial scales o variability

are enormous (Scheltema, 1986), evenwhen considering a single physical trans-port mechanism (see Box 1).

In contrast, v d pre ers tothe spread o larvae rom a spawningsource to a settlement site. This de ni-tion is consistent with the terrestrial lit-erature (natal dispersal in Clobert et al.,2001; Begon et al., 2006) that describesseed dispersal as the probability den-

sity unction o the number o seedsversus distance rom the adult source(i.e., the dispersal kernel) (Nathan andMuller-Landau, 2000; see Gerrodette,1981, or a rare marine example). Usingthe dispersal kernel, dispersal can beviewed as a probability that a releasedzygote will make it to settlement over

a certain distance, herein re erred toas dispersal distance. Larval transportis an important component o larvaldispersal, and broad dispersal requiressigni cant larval transport. Restricteddispersal, however, does not imply littlelarval transport (Figure 1). Further, pro-cesses and actors associated with theend o larval transport (i.e., settlement)also infuence dispersal, including settle-ment behavior, distribution o suitablesettlement sites, and re uge availability (Figure 2). Similarly, because spawninginitiates larval dispersal, spawning timeand location are important, as are actorsinfuencing spawning, including seasonand synchronicity o spawning, age andcondition o spawners, and ertiliza-tion success. In addition to the spatialdimensions inherent in larval transport,larval dispersal involves a survival prob-ability, and thus ood availability andpredation are important. The highestmortality in marine populations occurs

during the early li e stages, so mortal-

ity plays a large, but understudied,role in larval dispersal.

P p n c nn c vhas beende ned as the exchange o individualsamong geographically separated subpop-ulations (see Cowen et al., this issue) andis thought to be a key process or popu-lation replenishment, genetics, spread o

invasive species, and other phenomena(Cowen et al., 2006, this issue; Levin,2006). By this de nition, i the exchanis measured at the time o settlement,connectivity is essentially larval disper

rom one population to another (e.g.,Webster et al., 2002). Not all settlers wsurvive, however, and survival may beinfuenced by larval experience. Thus,connectivity is requently measured atsome point a ter settlement, once set-tlers survive to enter, or recruit to, the juvenile population. Functionally, how-ever, this point is somewhat arbitrary and di ers among taxa. A more precidemographic milestone is reproductionI settlers die without reproducing, dispersal is o questionable importance topopulation growth or spread o invasispecies. In this contribution we di ertiate between population connectivity,measured at the time o settlement, an

p d c v p p n c nn,de ned as the dispersal o individu-

als among subpopulations that survive

to reproduce. Reproductive populationconnectivity encompasses larval dis-persal but is also infuenced by post-settlement mortality (e.g., Hunt andScheibling, 1997; Doherty et al., 2004growth, and condition rom settlemento success ul reproduction. By the detion above, although dispersal o larva

t nd m n c n n p p nc nn c v d d

c p p n n v m n d p

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that do not survive to reproduce can play a role in population and community ecology, their contributions to reproduc-tive population connectivity are minimal(Figures 1 and 2).

larVal traNsPortr c n d n sc l v t n pThe term larval transport brings tomind small, passive larvae being movedthroughout the ocean by meso- andlarge-scale physical processes (Johnson,1939). This view has become a para-digmlarvae are released, trans-ported by mesoscale processes, mixedin a larval pool, and then randomly recruited to juvenile or adult habitat

(e.g., Roughgarden et al., 1988; Siegel etal., 2003). An increasing number o stud-ies, however, conclude that a signi cantamount o sel -recruitment occurs inmarine populations (Jones et al., 2005;Almany et al., 2007). These conclusionsare not in and o themselves surprising:a population is de ned as a sel -sustain-ing component o a species, and thussel -recruitment is a de ning attributeo a population (Sinclair, 1988). What issurprising is the relatively small spatialscales over which sel -recruitment hasbeen observed. For example, despite aplanktonic stage o 912 days, approxi-mately 30% o settling panda clown-

sh sel -recruited to an area o 0.5 km2

(Jones et al., 2005). The implication o

this and similar observations, combinedwith recent modeling and genetic studies(Cowen et al., 2000; Gerlach et al., 2007)

T m v m n v n n n b n x mp v p nd mp c nv v d n v n p . l v cc m

n c p p n c nv nc c c c v n pn n n b w m n , nd m c c nv nc

m w c nd w . on nd, w cmn fc n, n p n m n n, m d n n n b nd mp c v n p (P n d nd

lp z, 2002). a v n c , fc n nd n n b m d d b e N , n n nn p n m n n (Z mm m n nd

r b n, 1985). T , mp c v n ndv n p b n n d b n m c nd

mp c mp n n n d b vn d p c d nc d n p d c (~ 14.4 p d c c pp d w v ( w w k ; P n2002). in , v n p n nc md nc n n m n nd d k m ,

n c n b n.


Jess PiNeDa ([email protected]) is Associate Scientist, Department of Biology,

Woods Hole Oceanographic Institution,Woods Hole, MA, USA. JoNathaNa. hare is Research Marine Scientist,National Oceanic and Atmospheric

Administration, National Marine FisheriesService, Northeast Fisheries Service Center,

Narragansett Laboratory, Narragansett,RI, USA.su sPoNaugle is AssociateProfessor, Marine Biology and FisheriesDivision, Rosenstiel School of Marine and

Atmospheric Science, University of Miami,Miami, FL, USA.

BoX 1. VariaBility iN sPatial aND teMPoral sCales o larVal traNsPort

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1in c n b n w m n d c b ( ) w w w c nd b m ekm n n c , n l n z (1995), nd (b) z n , w c n nc d m d nd .

El Nio(several years)

Seasonalstratication(months)

Accumulation in internaltidal bore warm fronts

(seconds to hours)

and the constrained nearshore larval dis-tributions o littoral species (Barnett andJahn, 1987; Tapia and Pineda, 2007), isthat the spatial scales o larval transportmay be much smaller than previously recognized. These results indicate thatsmall-scale and nearshore physical pro-cesses play an important role in larvaltransport (Kings ord, 1990; Willis andOliver, 1990; Pineda, 1999).

N , C , ndoc n c C nFlows in nearshore, shallow environ-ments, including the sur zone, are di -

erent rom coastal and deep-ocean

fows mainly because o the shorelinebarrier, shallow depths, bathymetric

eatures associated with the continentalshel , and nearshore inputs o resh-water.1 Moreover, fows in nearshorewaters tend to be more complex thanin the deep and coastal ocean becausemany processes operate there, includ-ing sur ace gravity waves, buoyancy-driven fows, wind- orcing, sur ace andinternal tides, large-amplitude internalwaves and bores, and boundary-layere ects. These di erences between near-shore and coastal/open ocean hydrody-namics are important or larval trans-port. The shoreline barrier serves as a

topographic guide or coastally trappewaves and tends to steer fows in thealongshore direction (see Box 2). Tidaellipses that tend to be isomorphic inthe open ocean become compressednear the coast, and large-scale fows suas the Gul Stream and the HumboldtCurrent fow parallel to the shoreline,not perpendicular. Freshwater runo and large-scale currents running paral-lel to the coastline produce characterisstrati cation in the nearshore, such asshallowing o the thermocline near thcoastline in response to the Cali orniaCurrent (Hickey, 1979) and the FloridaCurrent/Gul Stream (Leaman et al.,

Oceanography s p mb 2007 2

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1989). Salinity (Thibaut et al., 1992)and water-column strati cation (Pinedaand Lpez, 2002) contribute to larvaltransport because sharper strati cationin shallow waters (e.g., Hickey, 1979)allows larvae o coastal species to exploit

vertically sheared fow to control hori-zontal distributions (Paris and Cowen,2004), and internal motions such asinternal tidal bores may transport larvaeonshore. Sur ace waves that break nearthe shore produce some mass transport,and storm systems that originate in thedeep ocean sometimes move onshore.Flows in the nearshore are broken by coastline topographic eatures such as

bays and capes, resulting in complexfows with smaller spatial coherence(see discussion in Okubo, 1994). This istrue or cross-shore coastal fows, whosecoherence scales are much smaller thanthe alongshore coastal fows (Brink,1999). The relative importance o theseprocesses varies with depth and distance

rom the shoreline (e.g., Lentz et al.,1999; Largier, 2003).

M d n NC s t n p bl sc P c

Clearly meso- and large-scale processesa ect larval transport, and most stud-ies emphasize these e ects. Large-scalephysical processes also infuence thesmaller-scale processes discussed above.Many large-scale circulation systemsand processes, such as eastern and west-ern boundary currents, El Nio, coastalupwelling, and coastally trapped waves,are energetic and coherent in the along-

shore direction, but can also modulatesmaller-scale processes in ways thatenhance or suppress larval transport.For example, as pointed out above, thestrength o the Cali ornia Current deter-mines the depth o the thermocline inshallow nearshore waters, with a stron-ger current resulting in a shallow ther-

mocline. A shallow thermocline createsvertically sheared environments that may restrict larval transport or species withdiel vertical migration; thus, interan-nual variability in the strength o theselarge-scale current systems might lead tovariability in dispersal, an untested spec-ulation. Consider the e ects o coastalupwelling, El Nio, and coastally trappedwaves on shallow water strati cationand cross-shore transport along the westcoasts o North and South America. Thecombination o strong coastal upwellingand El Nio produces weak nearshorestrati cation due to the upwelling o unstrati ed cold waters and the pilingup o mixed sur ace warm waters in thenearshore (Simpson, 1984; Zimmermanand Robertson, 1985). Both upwellingand El Nio result in decreased water-column strati cation, suppressing theshallowing o the thermocline by theinternal tide and the internal tidal bores,which, in turn, may result in decreasedonshore larval transport (recent work

o author Pineda and Manuel Lpez,Centro de Investigacin Cienti ca y deEducacin Superior de Ensenada). Incontrast, coastally trapped waves pro-duce a transient, small drop in sea levelthat is compensated by a large upli t-ing o the nearshore thermocline. Thisresults in the shallowing o the ther-mocline by the internal tide and larvaltransport by internal bore warm ronts

(Pineda and Lpez, 2002).

sm sc P c ndev n t p l v t n pSpatial and temporal scales are linkedin the ocean (Stommel, 1963), so theimportance o small-spatial-scale pro-cesses underscores the signi cance o

Time after spawning

D i s t a n c e f r o m s p a w n i n

g s i t e

x ,y0 0 0,t

x ,y ,t2 2 2

x ,y ,t1 1 1

x ,y ,t4 4 4 x ,y ,t4 4 5x ,y ,t3 3 3

larval transport

larval dispersal

reproductivepopulationconnectivity

s p a w n i n

g

d i s p e r s a

l

d i s t a n c e

s e t t l e m

e n t

r e p r o d u

c t i o n

1. r n p b w n p nd mp c mp n n v n p , v d p , nd p d c v p p n c nn c v p c . s v v p nd p c d. N m v n p d nc c n b n d p d

nc . W c c c n n p c w c d n x-y mt. a c n p c

xc p x , y0 nd x4 nd y4 , w c b n c. D nc c d b p n d n w d m nn ( . ., x , y c nd n x .)

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small-temporal-scale processes to larvaltransport. Moreover, meso- and large-scale processes can exhibit small-tem-poral-scale variability (Stommel, 1963)and be episodic (e.g., hurricanes). Larvalsettlement rom the plankton or many marine organisms is episodic, and it isnot uncommon to have the majority o a seasons settlement occur in a hand ulo days (Forward et al., 2004; Sponaugleet al., 2005). Even though settlementrecords imply transport events and areo ten correlated with various physi-cal actors, the observation o event-driven larval transport remains elusive.Similarly, larval distributions are o tenused to in er transport and the infu-ence o events (e.g., the occurrence o an eddy; Limouzy-Paris et al., 1997), but

ew studies have measured the move-ment o larvae in the water over time by event-type processes. When larval dis-tributions are sampled repeatedly overtime, they o er excellent views o theprocesses involved in larval transport

(Pepin and Helbig, 1997; Natunewiczand Epi anio, 2001), but due to samplinglimitations, such studies are rarely ableto observe the infuence o smaller-scaleprocesses. Examining the e ect o eventson transport is more straight orwardin a modeling contexta well-mod-eled example is the e ect o wind-driven events on settlement (Garvineet al., 1997; Brown et al., 2004)but

most circulation models do not cap-ture smaller-scale physical processes,

rontogenesis, rontal convergence anddivergence, intrusions, internal waves,and topographic e ects, particularly in the nearshore.

B v nd l v t n pAs our appreciation o small-scale physi-cal processes grows, so does our appre-ciation or the role o larval behavior in

infuencing larval transport. For many years, larvae were considered planktonic,that is, moving at the whim o oceancurrents but using eeding and preda-tor avoidance behaviors that resulted insmall-scale (millimeters to centimeters)movements (Blaxter, 1969). The viewo passive larvae gave way to the con-

cept that vertical swimming behavior,changes in buoyancy, and ontogeneticchanges in vertical position infuencethe horizontal movement o larvae; th

view was adopted early in estuarine ancoastal lagoon systems (Nelson, 1912;Pritchard, 1953; Bous eld, 1955) andlater in shel and open-ocean systems(Kelly et al., 1982; Cowen et al., 1993)Additionally, the infuence o larval setlement behavior on the speci c locatio settlement, at scales o meters to te

Connectivity = (larval dispersal, post-larval survival)

Larval behavior

Advection, diffusion

Dispersal = (larval transport, survival, spawning and settlement)

Larval transport = (physical transport, larval behavior)

2. T c nc p v n p , v d p , nd pn c nn c v . C w d n c c nc p .

n w n c nn c v b x m n d p nv v d n p p n c nn c v .

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l v n p n n nd p c n p n cnd n c mp n n ( . ., h ., 1999; M nd g ,

2005). T d nc n w c nv n n n c p c c np nd c nv n n b c c nd n d d n m c

p c v d n mp nd p c (W n n , 1983),d n p c p c d m n c nd n n p( . ., W n n nd B k v c , 1981), nd m m n m b nc n

w x cc n d b d n m ( . ., l n z ., 1999).a , p nk n p c v w d d n d m n n n w x(M n, 1993). B c n d n n w p p nd

c c v b n c d m n n, n p n x d p p n c n d b n v .

n p c w d v pm n m v pv w m , c n C n b n c

n p (t p nd P n d , 2007), c n p m cc p c , d v nd b w m ndm n n nd p d c . s m , p c m v p wn b v n m n b , c

a n c m n d n (Q n n ., 1999), v m m c j v n b . a c

mp z d n d v n p , bv p c p (h ., 1999), p cc nn c v . N nd c m n p p n

d n c , nd n m v m n v b w n p p n c n k p

p c d p p nc nn c d.

BoX 2. aloNg- aND Cross-shore PhysiCal tr aNsPort ProCesses

o meters, was recognized as important(e.g., Crisp, 1976; Raimondi, 1991).

More recent research shows thatlarvae also have horizontal swimmingcapabilities that improve with develop-ment (see review by Leis, 2006). Forexample, larvae o a damsel sh swamcontinuously or 39 hours without ood,covering a distance equivalent to 19 km(Stobutzki, 1997). Similarly, larval lob-sters and early pelagic stages o cepha-lopods are good swimmers (Villanuevaet al., 1996; Je s and Holland, 2000).In combination with the capability toswim vertically and horizontally, larvaeo both invertebrates and vertebratescan orient and potentially navigateover short (meter-to-kilometer) to long(10-to-100-km) distances, using light,

sound, smell, and possibly magnetism,electric elds, and wave swell (e.g.,Kings ord et al., 2002; Gerlach et al.,2007). Clearly, larvae are complex andcapable organisms that develop the abil-ity to eed, avoid predation, and movewithin the pelagic environment. Thus, inthe equation o larval transport, behav-ior plays an equally important role asadvection and di usion.

larVal traNsPort:researCh NeeDsid n c n N l vt n p M c n mKnowledge o larval transport in near-shore environments is very limited.Major drawbacks include lack o rigor-ous knowledge o the suspected physical

mechanisms involved in larval transport,and ignorance o other potential trans-port mechanisms (see Cowen, 2002, or areview). Physical mechanisms that coulda ect transport include sur ace grav-ity waves (Monismith and Fong, 2004),submeso- and mesoscale eddies (Bassinet al., 2005; Sponaugle et al., 2005), baro-tropic tidal currents (Hare et al., 2005;Queiroga et al., 2006), and cross-shorewinds (Tapia et al., 2004).

Some proposed mechanisms have notbeen tested rigorously in eld condi-tions. Moreover, the logistical di culty o studying transport sometimes canpush researchers to use weak in eren-tial approaches, such as in erring larvaltransport mechanisms rom settlementdata (Pineda, 2000; Queiroga et al.,

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in larger-scale models, thereby capturingthe large-scale aspects o larval trans-port, the modulation o small-scale pro-cesses by large-scale orcing, and the very small-scale processes (e.g., turbulence)where larval swimming capabilitiesand behavior become overly important(see discussion in Metaxas, 2001). Evenmodeling a single, relatively straight-

orward process, such as the accumula-tion o particles in gravity currents, canbe extremely complex (e.g., Scotti andPineda, 2007). Thus, using numericalmodels or in erring larval transportwhen poorly studied processes may beimportant, or where the physical orcingis unknown, is dire. On the other hand, itis clear that numerical models are pow-er ul tools in settings where processesare well known and in cases where eldhydrodynamics are well simulated by themodel (e.g., Reyns et al., 2006). Thus, wesuggest that bottlenecks in understand-ing larval transport are less related tonumerical modeling than to the mecha-

nistic knowledge o larval transport.

C n ad p v s mp nIt is unclear how much larval transportoccurs during episodic events and howmuch occurs during mean condi-tions. Sharp peaks in settlement timeseries and studies o larval transportby wind and internal motions suggestthat transport can be sporadic, larvae

extremely patchy, or both (see Pineda,2000, or discussion). Time-series mea-surements o relevant hydrodynamicsand larval distributions during larvaltransport are o limited use when mea-surements cannot be taken with the nec-essary requency and spatial resolutionto describe the processes with su cient

detail. Furthermore, surveys by researcvessels diligently planned in advancedo not guarantee that larval-transportevents will happen during the surveys.Adaptive sampling, de ned as samplinin response to an event, is a solution tothese dilemmas; it has been used suc-cess ully to sample hydrodynamics anlarval distributions during transportby internal tidal bores (Pineda, 1994,1999). Adaptive sampling is challenginhowever, because it is hypothesis basesampling is initiated in response to areal-time change in a time-dependentvariable, such as temperature or winddirection, that is integral to the hypothesized larval transport mechanism.Adaptive sampling is there ore a stringent hypothesis test, because i larvaltransport does not occur as expected, thypothesis is rejected. Adaptive samplis also logistically di cult. I the eveare sporadic, and the sampling is ship-board, adaptive sampling requires hav-ing a vessel and crew on standby ready

to sample or long periods, an expen-sive prospect or anxious researchers.Conceivably, remote sampling systemsinitiated in response to events could beconstructed with o -the-shel gear anew technologies currently under deveopment such as in situ molecular detection o larvae (e.g., Go redi et al., 2Thus, similar to the limitations in modeling larval transport, adaptive samplin

is limited in part by technology and inpart by the development o testable,mechanistic hypotheses.

B k n B v BThe incorporation o larval behavior

ully into the larval transport equationrequires several important advances.

2006). The lure o mesoscale processesand satellite oceanography has provedirresistible or some shallow-waterecologists, resulting in an overempha-sis on explanations based on mesoscaleprocesses while disregarding nearshoreprocesses and mechanisms that cannotbe studied remotely. Unambiguous iden-ti cation o the mechanisms o larvaltransport is rare, and testing alternativeexplanations is almost unheard o . Thus,there is a serious need to ollow up someo these weakly ounded hypotheses withrigorous tests. With limited knowledgeo nearshore larval transport, it seemsthat assessing the relative contributionso various physical transport mecha-nisms in larval transport or a given casestudy is, so ar, only a utopian hope. The

eld will be mature when such a study can be proposed and accomplished.

Understanding the role o small-scaleprocesses in larval transport is also lim-ited by modeling capabilities. Large-scaleand mesoscale models orced by winds

and the sur ace tide are now common-place (see Werner et al., this issue). Thespatial resolution o these models isincreasing and extending into nearshoreareas (e.g., Chen et al., 2006). Decreasedgrid size, however, is only one aspect o resolving smaller-scale processes. Small-scale processes, such as sur ace waves,internal waves, and propagating conver-gences, need to be included. Currently,

no numerical model appears capableo simultaneously resolving Lagrangiantransport caused by, or example, shal-lowing internal tides, sea breeze, large-amplitude internal waves, and sur-

ace gravity waves. Further, accurately modeling larval transport will requireembedding these small-scale processes

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First, hypotheses on the role o behaviorin transport need to be developed andtested. Colby (1988) argued that passiveadvection and di usionshould be the nullhypothesis or studies o larval transport.In an early example o this approach,Woods and Hargis (1971) comparedthe distribution o coal particles withthat o similarly sized oyster larvae andconcluded that larvae were not beingtransported passively. A study on ascid-ian tadpole larvae ound that dispersaldistance was shorter in swimming lar-vae than in nonswimming individualso similar size and shape (Bingham andYoung, 1991). Similarly, Arnold et al.(2005) ollowed a cohort o larval hardclams and ound their distribution di -

ered rom dye distributions and rommodeled distributions based on passiveparticles. There are other examples o the use o a hypothesis-testing approach

or evaluating the processes that a ectlarval transport (e.g., Hare et al., 2002).This approach should be expanded to

take advantage o advances in modelingas well as in eld and laboratory studies.Behavioral hypotheses rom laboratory studies are attractive because quanti ca-tion o hydrodynamics and behavior is

easible, but these hypotheses should betested in eld conditions, and vice versa.

Second, the incorporation o behav-iors into models o transport needs to berule-based rather than deterministic, and

individual variability should be consid-ered. Most transport models that includelarval behavior use population-leveldescriptions o distributions or swim-ming speeds and apply them to particlesreleased in the model (Hare et al., 1999).Another approach is to provide a set o behavioral rules that attempt to capture

the trade-o s between eeding and pre-dation; these rules result in vertical (andpotentially horizontal) responses to vari-ous cues (Titelman and Fiksen, 2004;Fiksen et al., in press). Although theimportance o time-dependent behav-iors, such as diel, tidal, and ontogenetic,is well recognized, little is known aboutadaptive behavior on scales o secondsto minutes, where larvae might respondto transient physical and biological

eatures. We know that larvae respondbehaviorally to a number o actors, suchas time o day, light, temperature, tur-bulence, pressure, and ood availability,and that some o these responses infu-ence transport, but only a ew behaviors

acilitating transport have been identi-ed (e.g., Boehlert and Mundy, 1988;

DiBacco et al., 2001). For example, eldobservations, modeling, and labora-tory experiments imply that swimmingup behaviors in response to transientdownwelling fows in propagating ea-tures determine e cient larval transport

(Pineda, 1999; Scotti and Pineda, 2007).To incorporate our understanding o behavior into rule-based models willrequire a hypothesis-based approach.Without hypotheses, we run the risk o evaluating the e ect o multiple irrel-evant behavioral scenarios on larvaltransport. This rule-based approachcoupled with more studies on adaptivebehavior and well-developed biophysi-

cal, individual-based models (e.g., Loughet al., 2005, and recent observations o Claudio DiBacco o Bed ord Instituteo Oceanography, author Pineda, andKarl Hel rich o WHOI), will greatly advance our understanding o the com-bined roles o advection, di usion,and larval behavior.

Third, most research has ocused onhow larval behavior a ects advection,but the infuence o behavior on di u-sion requires more emphasis. Using anadvection-di usion-mortality model,Cowen et al. (2000) estimate that suc-cess ul larval transport to coral ree habi-tats diminishes sharply when di usionrates increase rom 0 to 100 m2 s-1 (thelatter is a typical di usion rate used inlarval transport studies; see also Okubo,1994). However, the assumption thatlarvae di use passively in the marineenvironment likely does not hold, par-ticularly or older larval stages. Peaksin settlement must result rom high-density patches o larvae reaching adulthabitats, and these coherent patchesrun counter to hypothesized di usion.Natunewicz and Epi anio (2001) ol-lowed discrete patches o crab larvae

or up to six days and hypothesized thatassociative swimming behaviors mightbe responsible or patch maintenance.A U-shaped patchiness-at-age unction

has been described or the larval stageso several sh species, and this shape hasbeen interpreted as initial di usion withsubsequent schooling (Matsuura andHewitt, 1995). In addition, larvae may remain in thin layers o ood (Lasker,1975) and reduce their di usion owingto vertical di erences in fow (shear di -

usion). Larvae can also accumulate atupwelling and downwelling ronts by

swimming into the current (e.g., Franks,1992; Metaxas, 2001), thereby reducingdi usion. Thus, small-scale vertical andhorizontal larval behavioral responsesmay limit di usion and greatly a ectlarval transport. Consequently, the useo advection-di usion models to under-stand larval transport requires great

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care. For example, Hill (1991) under-scored the limitations o an advection-di usion-mortality model in cases whenactive vertical positioning o larvae wasexpected, and Okubo (1994) warned thata horizontal di usive model would notwork in settings with strong convergentfows, a widespread phenomenon incoastal and nearshore settings.

larVal DisPersalD n n D p K nMost attempts to describe dispersal ker-nels have emphasized larval transport(e.g., Bots ord et al., 1994), but otherprocesses such as spawning, settlement,pelagic larval duration, and survival alsoinfuence larval dispersal (Edwards et al.,in press). Many marine species releasetheir o spring at speci c locations andtimes, using speci c behaviors. Forexample, relatively sedentary blueheadwrasse spawn daily at particular ree spawning sites that have been used or years (Warner, 1988). Similarly, several

sh species spawn in circular motionsthat may create hydrodynamic vortexes(Okubo, 1988; Heyman et al., 2005). Theinfuence o these small-scale events onlarval dispersal over periods o weeks isunknown. On a larger scale, a number o motile species, including snappers, her-ring, and blue crabs, move to particularlocations or spawning (Carr et al., 2004;Heyman et al., 2005). In the temporal

domain, many coral species participatein annual mass spawning events, withmore than 60% o species spawning overthe course o several days (Babcock etal., 1994), and crabs and barnacles tendto release their larvae at certain phaseso the tide or the day (Morgan, 1995;Macho et al., 2005). While such spawn-

ing behaviors have long been thought tomaximize larval survival (e.g., Hughes etal., 2000), the overall e ect o localizedand punctuated spawning on larval dis-persal is unclear.

Moreover, where individuals end theirplanktonic stage is also an important

component o larval dispersal. Larvaldurations o some species are xedwhile others are fexible (Pechenik, 1986;Cowen, 1991). Some species have very narrow habitat requirements or the con-tinuation o the li e cycle, such as rivermouths on isolated oceanic islands orsome gobies, wave-beaten rocky points

or gooseneck barnacles, and speci cspecies o anemones or some ree sh(Radtke et al., 1988; Cruz, 2000; Joneset al., 2005). Other species have broadhabitat requirements such as eurytopicPachygrapsus crabs (Hiatt, 1948) andfounders o the genusEtropus (Walsh etal., 2006). For most species, only a subseto locations will support the continu-ation o the li e cycle; these locations

must be reached within the time windowo possible settlement. Understandingthese habitat and time constraints willbe necessary to observe and model dis-persal kernels. A number o models haveincluded such considerations at a rela-tively large scale, or example, assum-ing modeled larvae that arrive within

1015 km o known habitat have suc-cess ully settled (Hare et al., 1999; Paet al., 2005). How larvae transverse thelast 10 km is unknown largely becausethe exclusion o smaller-scale processin models and the inability to includerealistic behaviors (see above).

The dispersal kernel also is dependeon larval mortality. Most studies o lardispersal, however, either do not con-sider larval mortality (Hare et al., 1999consider spatially homogenous mortality (Cowen et al., 2000), or assume lowmortality (Gaylord and Gaines, 2000).Modeling studies that assume low mor

talities should be reconsidered in lighto observed higher mortalities (e.g.,Rumrill, 1990); use o high mortalitiein dispersal models requently yieldslower maximum dispersal estimates ththose obtained assuming low mortality(Cowen et al., 2000; Ellien et al., 2004Tapia and Pineda, 2007). Di erentialsurvival o larvae during transport contributes to de ning the dispersal kerne

in potentially numerous species-speciways. The ecological literature is richwith examples and models in which throle o spatial heterogeneity in mortality shapes subsequent patterns in abundance, distribution, and demographics.These concepts, however, have yet to bapplied to mortality in pelagic early li

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stages. It is also clear that not all larvaeare equal, and the range o traits willresult in selective survival (see later sec-tion on Population Connectivity).

Larval duration also infuences sur-vival probability. Pelagic larval dura-tion (PLD) must be correlated with thedispersal kernel or the simple reasonthat species with short PLD must havereduced larval transport and relatively short dispersal kernels; PLD is aconstraining variable or dispersal. Incontrast, long PLDs do not necessarily yield broad dispersal kernels, as larvalbehavior breaks the direct-proportionalrelationship between PLD and dispersaldistance, both or sh and invertebrates(Sponaugle et al., 2002). O course, longPLD yields higher cumulative mortali-ties than short PLD when everything

else is equal (i.e., same daily mortality or species with short and long PLD; see

Hare and Cowen, 1997). It is also unclearhow variables infuencing PLD, such astemperature and ood (Scheltema andWilliams, 1982), may infuence the dis-persal kernel (see OConnor et al., 2007,

or model predictions). Thus, the rela-tionship between PLD and dispersal isambiguous except or species with very short larval durations (see discussion inSponaugle et al., 2002).

D p e m n C oc nGiven the complexity o larval dispersal,it is not surprising that measurement o a dispersal kernel in the marine environ-ment is extraordinarily rare (Shanks etal., 2003). Gerrodette (1981) measured

the dispersal o planula larvae romadults in a temperate solitary coral and

ound that mean dispersal distance romthe parent was < 50 cm. Similar workwith ascidians quanti ed dispersal romspawning to settlement, but the pelagicstage o ascidians is short (hours), larvaeare large (millimeters), and mortality islow (< 90%) (Olson and McPherson,1987), making it possible to ollow indi-viduals rom the beginning to the end o the pelagic stage (see also Bingham andYoung, 1991). Work on an isolated ree indicated that most acroporid and pocil-loporid corals recruited in experimentalmoorings within 300 m rom the ree ,and that spat mortality decreased withdistance rom the ree (Sammarco andAndrews, 1989). Several studies ollowedpatches o more typical marine larvae


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(PLD o weeks, size < 110 mm, andhigh mortality), but these e orts are nottrue measures o larval dispersal becausethe spawning and ending locationswere in erred (Pepin and Helbig, 1997;Natunewicz and Epi anio, 2001; Parisand Cowen, 2004). Other studies markedspawned eggs and then collected o -spring at the end o their planktonic stage(Jones et al., 2005; Almany et al., 2007);these studies provide a partial measure,but not a complete description, o thedispersal kernel because all potentialending locations could not be sampled.Although dispersal kernels will eventually be ully quanti ed or some species insome systems, the measurement o theseprobability distributions in the marineenvironment will remain extremely rare.

It is easier to obtain dispersal kernelswith models than with eld measure-ments. Some models consider simpli-

ed situations using advection-di usionmodels. More complex numerical circu-lation models coupled with Lagrangian

particle-tracking algorithms ollowparticles released at multiple locationsand multiple times and have proveninstrumental in estimating dispersal ker-nels in the marine environment (Cowenet al., 2000; see also Werner et al., thisissue). Edwards et al. (in press) used a

ully orthogonal approach to examinethe e ects o di erent actors on generictwo-dimensional dispersal kernels esti-

mated rom a three-dimensional circu-lation model o the Southeast UnitedStates shel . This study ound that timeand place o initial release were mostimportant in determining the position o the dispersal kernel, and that dispersionand PLD were most critical in determin-ing the spread o the dispersal kernel.

Larval behavior was not as important,but horizontal swimming behavior wasnot included and depth-strati ed cur-rents were minimal through most o themodeling domain, limiting the e ect o di erent vertical positions.

larVal DisPersal:researCh NeeDs

d ob v n D pThe paradigm o broad dispersal o

sh and invertebrate larvae is givingway to the notion o restricted disper-sal, mainly because o studies nd-ing: (1) unexpected high levels o sel -recruitment, (2) high larval mortality rates, and (3) restricted scales o larvaltransport (see above). Still, the domi-nant scales o dispersal are not known.Solid empirical estimates o dispersalare needed to guide eld and numeri-cal modeling studies to address ques-tions such as: What regions o the oceanshould researchers ocus on? What pro-cesses must be included in the models?

Studying dispersal is challenging, and orsh and invertebrate species with long

and typical larval durations (i.e., aboutour weeks or temperate invertebrates;

Levin and Bridges, 1995), knowledge willbe gained incrementally by using mul-tiple approaches, including: (1) empiricalestimates o larval origin, such as natu-ral and arti cial tags and genetic dis-tance and structure, (2) a mechanistic

understanding o larval transport,(3) assessment o how the space and timeo spawning and settlement infuencedispersal, (4) trophodynamic studies toaddress the infuence o pelagic patchi-ness and structure on the larval jour-ney rom spawning to settlement, and(5) improved mortality estimates in dis-

persal models in locations where physiprocesses are well known.

When empirical estimates o dispersal are obtained, it is crucial that they bused to test the assumptions and hypoteses resulting rom both simple andcomplex models. Robust measuremento dispersal will be rare and opportunties to evaluate and test models must nbe lost. In this way, the skill o modelscan be assessed and improved throughan iterative process o observation andmodeling, and the resulting dispersalkernels can be part o larger studies oconnectivity with increasing con dencAlthough the challenges are immense, emphasize that solid empirical estimato dispersal are necessary to guide uther eld studies and numerical modeling; theoretical developments and modeling o spatial population processes aconnectivity may be utile unless we gmore observationally based knowledgeo larval dispersal.

PoPulatioN CoNNeCtiVityt C nc p P p nC nn c vA mechanistic understanding o larvaldispersal is su cient or determiningpopulation connectivity at time o settment. Knowledge o population con-nectivity at the time o settlement orshortly therea ter may be adequate orsome objectives because subadult indi-

viduals use resources, interact with aduand other members o the community and in some instances, sustain sher-ies.Reproductive population connectiv-ity, on the other hand, is the exchange oindividuals that eventually reproduce.Accordingly, or benthic marine specieit is not only a unction o larval dispe

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(including survivorship o larvae duringtransit), but also o post-settlement and juvenile survival to the point o repro-duction (Figures 1 and 2). Reproductivepopulation connectivity can be expressedas the number o individuals rom site aand population A that disperse to site bcontaining population B and reproducethere per unit time. Thus, during devel-opment to the adult stage (which var-ies greatly among species, rom days tomultiple years), juveniles must survive,grow, mature, and reproduce. As charac-teristics o settlers are o ten variable andthose surviving to reproduce may not bea random sample o the settlers, simply tracking larval trajectories rom spawn-ing to settlement is insu cient to quan-ti y reproductive population connectivity.The remainder o this discussion consid-ers the ecological processes contributingto reproductive population connectivity.

For a population to be ecologically sustained, a minimum number o o -spring must mature and reproduce over

time intervals dictated by species longev-ity. Identi ying this number is essentialto parameterize population models, butan equally important consideration is thecomposition o the survivors that makeup this number: What are the character-istics o dispersers that lead to success ulrecruitment? Which o those recruits willthen survive to reproduce? Recent evi-dence points to important infuences o

spawning patterns, maternal e ects, andpelagic experience on larval size, growth,condition, and survival. Furthermore,many o these larval traits carry overand infuence juvenile survival. However,comparatively little is known about thelinkages between these early li e phenom-ena and adult survival and reproduction.

V n n l v t nds v v D n P c sMost larvae exhibit variation in early li e history (ELH) traits, such as size ata given age and growth rate. This varia-tion can be introduced as early as the eggstage, when di erential size, age, condi-tion, or stress level o the mother caninfuence quality o the spawned eggs(Berkeley et al., 2004; McCormick, 2006).Larval encounter with variable pelagicenvironments also infuences larvalgrowth and survival. Water temperatureplays a central role in regulating metabo-lism and growth (Houde, 1989), withlarvae in di erent temperatures exhibit-ing variable ELH traits (Meekan et al.,2003; Sponaugle et al., 2006). Sustainedgrowth requires adequate ood; there-

ore, variable access to ood also a ectslarval traits and survival. Transit acrossnutrient-poor open oceans may be par-ticularly di cult or species with highgrowth rates. Access to ood and avoid-ance o predation or other develop-

mental conditions may be related to thetiming o spawning, such that particularwindows o time result in higher larvalsurvivorship (Cushing, 1990; Baumannet al., 2006). Encounter with oceano-graphic eatures such as ronts or meso-scale eddies can also infuence ood sup-ply and exposure to predators (Grimesand Kings ord, 1996; Sponaugle andPinkard, 2004). Thus, a complex oceano-

graphic environment coupled withvariable egg quality at spawning resultsin a pool o larvae with variable traits(Jarrett, 2003; Lee et al., 2006; Sponaugleand Grorud-Colvert, 2006).

Survival o pelagic larvae is typically nonrandom and proceeds according tothree general concepts o the growth-

mortality hypotheses (reviewed inAnderson, 1988). Theoretically, survivorsshould be those larvae that are larger at agiven age (bigger is better hypothesis;Miller et al., 1988), grow aster (growth-rate hypothesis; Bailey and Houde,1989), and/or move through an early stage more rapidly (stage-durationhypothesis; Anderson, 1988). Larvae o a diversity o marine sh (e.g., Meekanand Fortier, 1996; Hare and Cowen,1997; Meekan et al., 2006) appear toadhere (to varying degrees) to aspects o these overarching concepts. Di erentialsurvival o larvae due to their pelagicexperience and ELH traits can infu-ence the magnitude o larval settlementpulses. Variation in the magnitude o settlement events has been related tovariable larval growth throughout orduring particular periods o larval li e(e.g., Bergenius et al., 2002; Jenkins andKing, 2006; Sponaugle et al., 2006)

in nc l v t n

J v n s v vSettlement o larvae to the benthos isa risky event plagued with high levelso predation mortality (e.g., Hunt andScheibling, 1997; Doherty et al., 2004);thus, additional selective loss typically occurs during this period. Most marinespecies undergo a metamorphosisbetween the larval and juvenile stages asthey move between radically di erent

environments. While metamorphosisenables closer adaptation to stage-spe-ci c environments (Wilbur, 1980), larvalhistory is not erased and accompaniesthis transition (Pechenik et al., 1998).Importantly, recent studies have begunlinking these two stages and investigatinghow larval traits infuence juvenile sur-

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vival. Traits exhibited by settling larvaeas a consequence o pelagic constraintsand selective pressures have the potentialto carry over and infuence survival o juveniles. For example, larval growth,size, and condition infuence the survi-vorship o juvenile sponges, molluscs,barnacles, bryozoans, and shes (e.g.,Searcy and Sponaugle, 2001; Pechenik etal., 2002; Jarrett, 2003; McCormick andHoey, 2004; Phillips, 2004; Marshall etal., 2006; Sponaugle and Grorud-Colvert,2006). The potential exists or sometraits that are advantageous to larvaeto become subsequently detrimental to juveniles or vice versa. For example, crabzoeae reared at reduced salinities su erhigher mortality as larvae, but metamor-phose into larger juveniles (Gimnez andAnger, 2003), and a short pelagic larvalduration enables sh larvae to escape thepredation in the plankton, but resultsin smaller settlers (e.g., Sponaugle et al.2006), which in some cases may be moresusceptible to predation (Anderson,

1988). Most studies have ocused on con-sequences to juveniles and somewhat lesson the trade o s associated with confict-ing constraints in complex li e histories.

s v v p B nd J v n sAlthough events during larval li e canplay an important role in early juve-nile survival, much less is known about

how these traits are carried through orlost rom individuals that survive toreproduce. Studies on larval dispersal orpopulation connectivity typically de nerecruitment as entry into the juvenilepopulation, not to the adult popula-tion. Thus, settlers are tracked at most tothe point o settlement or through the

rst ew days or weeks as juveniles. Weknow little about the settlers that eventu-ally survive to reproduce. It is generally substantially more time-consuming andlogistically challenging to track cohortso settlers all the way to reproduction. A

ew recent studies have had some successollowing species that mature rapidly.

Pineda et al. (2006) sampled barnaclesthat settled over an 89-day period untilthey reproduced 11 months later and

ound that survivors settled during anarrow 21-day recruitment window.Meekan et al. (2006) tracked a singlecohort o a ast-growing coastal sh and

ound that despite strong selective loss

during early stages, there was no addi-tional selective mortality between the juvenile and adult stages. For bryozoansin an experimental manipulation, how-ever, adults that were larger as larvaehad higher survival rates and producedlarger larvae themselves than those thatwere smaller as larvae, although delayingmetamorphosis erased this relationship(Marshall and Keough, 2006). Optimal

traits may vary with the environmentencountered by the larval, juvenile, oradult stages, as evident or a snail (Moranand Emlet, 2001) and colonial ascid-ian (Marshall et al., 2006). Thus, traitsobtained during early stages have thepotential or long-term e ects on laterstages, but many complex interrelation-

ships likely infuence the outcome. Whecarryover e ects occur, they may perbecome ampli ed, or, instead, be com-pensated or during subsequent stages(Podolsky and Moran, 2006). In short,simply reaching a settlement site doesnot guarantee that larvae will possess tnecessary traits to survive to reproduce

PoPulatioN CoNNeCtiVity:researCh NeeDsThe undamental challenge in popula-tion connectivity studies is to determinthe source populations o settling larvand the settlement sites o dispersing lvae. In short, all the research needs ideti ed under the larval transport and dis

persal sections sum together as researcneeds or population connectivity. Inaddition, there is a need to link materne ects and larval processes to early jnile survival and, in the case o repro-ductive population connectivity, to thepoint o reproduction. Because repro-ductive population connectivity per seis de ned as the exchange o individuthat eventually reproduce, tracking dis

persing larvae to the point o settlemeor juvenile recruitment, while importan

or some purposes, is unctionally inscient. New e orts to track settlers to

reproduction will initially advance witshorter-lived sessile species. Eventualllong-term, labor-intensive studies willbe needed to increase our understandin

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o reproductive population connectivity o longer-lived mobile species. There isa rich history o marine ecological workexamining the relative importance o recruitment versus density-dependent,post-settlement processes in structuringbenthic populations (Caley et al., 1996),but we need to move beyond numeri-

cal responses and re ne the question toocus on trait-based ecological linkages

among all stages. Real measures o repro-ductive population connectivity requirean understanding o who is surviving toreproduce and why.

As there is ample evidence that larvalgrowth and condition can infuence per-

ormance in later stages, rom a practi-cal point o view we need more reliablemeasures o condition. The coarsestmeasures o condition o ten use size asa proxy (e.g., many invertebrates), whileothers measure organic (Jarrett, 2003)or lipid content (Hentschel and Emlet,2000), RNA/DNA ratios (Suthers et al.,1996; Lee et al., 2006), or ( or shes)otolith-based measures (e.g., Sponaugle

et al., 2006), all o which have some limi-tations. As new genomic techniques aredeveloped, perhaps new measures o per-

ormance can be incorporated into bothobservational and manipulative studies.

Finally, ocusing on the individu-als that survive to reproduce may guidelarval transport and dispersal studies;

i settlers that survive to reproductionare only spawned at timet and site x , y ,or i success ul individuals only settlein recruitment windows coincidingwith physical-transport processes p and

eeding and prey environmentse, thevast parameter space that potentially a ects pelagic eggs and larvae, and vexes

researchers, may be e ectively reduced toa more manageable set.

aCKNoWleDgeMeNtsWe thank the National Oceanic andAtmospheric Administration, theNational Science Foundation, and theWoods Hole Oceanographic Institution

or supporting our work, and JohnManderson, David Mountain, NathalieReyns, Vicke Starczak, Fabin Tapia,Simon Thorrold, and an anonymousreviewer or constructive criticisms.

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