Patterns of settlement within a restored mussel bed site

The failure of depleted bivalve populations to recover naturally is often due to a breakdown in recruitment processes, such as a lack of larvae or suitable settlement substrate. The identification of these recruitment limitations are a critical initial step for determining an appropriate strategy for restoration. A lack of larval supply and settlement substrate have both been suggested as possible causes of the failure of natural recovery in the green‐lipped mussel population in the Hauraki Gulf, New Zealand, and were therefore examined in this study. Larval mussels settled throughout most of the year directly onto artificial collectors placed within restored adult mussel beds and in the immediate vicinity of the beds at comparatively lower levels than for remnant populations throughout New Zealand. This low settlement coupled with a lack of recruitment to the restored mussel beds suggests larval supply may be limited in the Hauraki Gulf and warrants further examinations into larval dispersal and retention. Larval settlement was also higher on collectors within mussel beds, suggesting the presence of adults enhances larval settlement and highlights the importance of transplanting adult mussels for reestablishing mussel populations.     

Habitat selection as a source of inter‐specific differences in recruitment of two diadromous fish species

• 1 For aquatic species with highly dispersive offspring, the addition of new individuals into an area (recruitment) is a key process in determining local population size so understanding the causes of recruitment variability is critical. While three general causative mechanisms have been identified (the supply of individuals, habitat selection and mortality), we have a limited understanding of how variation in each is generated, and the consequences this may have for the spatial and temporal distribution of recruits.
• 2 We examined whether active habitat selection during settlement could be the cause of variability in populations of two diadromous fish species using a field survey and laboratory‐based choice experiments. If larval behaviour is important, we predicted there would be inter‐specific differences in abundance between sites during the survey, and that larvae would prefer water collected from sites with higher conspecific abundances during the experiments.
• 3 During the field survey, significant differences were detected between two rivers (the Cumberland and Grey), with one species (Galaxias maculatus) found in higher abundances at one site (the Cumberland River) while comparable numbers of a closely related species (Galaxias brevipinnis) were caught at both sites. Laboratory choice experiments were conducted to determine whether larval preferences during settlement could be the cause of these differences. G. maculatus larvae showed a preference for freshwater over saltwater, indicating that the fish may be responding to reduced salinities around river mouths during settlement. The results of a second experiment were consistent with the notion that larval preferences could be the mechanism driving differences in the populations of the two rivers, with G. maculatus preferring water collected from the Cumberland River while G. brevipinnis did not prefer water from either river.
• 4 These results demonstrate that active habitat selection may be important in establishing spatial patterns of larvae at settlement, and that multiple cues are likely to be involved. This study also demonstrates that the behaviours exhibited by individuals can strongly influence the structure and dynamics of populations of aquatic species with complex life cycles.

     

Habitat specialization and adaptive phenotypic divergence of anuran populations

We tested for adaptive population structure in the frog Rana temporaria by rearing tadpoles from 23 populations in a common garden experiment, with and without larval dragonfly predators. The goal was to compare tadpole phenotypes with the habitats of their source ponds. The choice of traits and habitat variables was guided by prior information about phenotypic function. There were large differences among populations in life history, behaviour, morphological shape, and the predator-induced plasticities in most of these. Body size and behaviour were correlated with predation risk in the source pond, in agreement with adaptive population divergence. Tadpoles from large sunny ponds were morphologically distinct from those inhabiting small woodland ponds, although here an adaptive explanation was unclear. There was no evidence that plasticity evolves in populations exposed to more variable environments. Much among-population variation in phenotype and plasticity was not associated with habitat, perhaps reflecting rapid changes in wetland habitats.     

Epizootiology of Amblyospora stimuli (Microsporidiida: Amblyosporidae) infections in field populations of a univoltine mosquito, Aedes stimulans (Diptera: Culicidae), inhabiting a temporary vernal pool.

The epizootiology of the microsporidium Amblyospora stimuli was studied in natural populations of a univoltine mosquito, Aedes stimulans, inhabiting a temporary vernal pool over an 18-year period. The yearly prevalence of benign oenocytic infections in adult females was variable, ranging from 1.0 to 9.6% (mean = 5.1%). The yearly prevalence of transovarially transmitted meiospore infections in larval populations was consistently lower but less variable, ranging from 1.3 to 5.9% (mean = 3.5%). Meiospore infections in F(1)-generation larvae were significantly correlated with infections in parental-generation females, thus suggesting that larval infection rates could be substantially increased if methods were available to facilitate transmission of A. stimuli to a larger portion of the female population via inundative or inoculative release of infected copepods. No correlation was found when infections in filial-generation adult females were measured against meiospore infections in larvae from the preceding year. Analysis of yearly prevalence data using Fine's Fundamental Vertical Transmission Equation revealed low rates of horizontal transmission from the intermediate copepod host to female larvae in most years, ranging from 0.1 to 8.7% (mean = 3.1%). A. stimuli is enzootic, persists at a very low level, and has minimal impact on Ae. stimulans populations at this site. The low incidence rate of horizontal transmission to larvae appears to be due largely to a paucity of copepods and is a major factor that limits the abundance and subsequent proliferation of A. stimuli in Ae. stimulans populations at this locale. Results support the view that host-parasite cospeciation is an important mechanism of evolution in this group of mosquito/copepod microsporidia.     

Reproductive output and duration of the pelagic larval stage determine seascape-wide connectivity of marine populations

Connectivity among marine populations is critical for persistence of metapopulations, coping with climate change, and determining the geographic distribution of species. The influence of pelagic larval duration (PLD) on connectivity has been studied extensively, but relatively little is known about the influence of other biological parameters, such as the survival and behavior of larvae, and the fecundity of adults, on population connectivity. Furthermore, the interaction between the seascape (habitat structure and currents) and these biological parameters is unclear. We explore these interactions using a biophysical model of larval dispersal across the Indo-Pacific. We describe an approach that quantifies geographic patterns of connectivity from demographically relevant to evolutionarily significant levels across a range of species. We predict that at least 95% of larval settlement occurs within 155?km of the source population and within 13 days irrespective of the species' life history, yet long-distant connections remain likely. Self-recruitment is primarily driven by the local oceanography, larval mortality, and the larval precompetency period, whereas broad-scale connectivity is strongly influenced by reproductive output (abundance and fecundity of adults) and the length of PLD. The networks we have created are geographically explicit models of marine connectivity that define dispersal corridors, barriers, and the emergent structure of marine populations. These models provide hypotheses for empirical testing.     

Breedng season, larval development and diseprsal of Lingula anatina (Brachiopoda, Inarticulata) from Townsville, Australia

A discrete but protracted breeding season, extending from October to April, was defined for Lingula anatina from Townsville, Australia, using gonad indices, sizes of developing ova, occurrence of planktonic larvae and analysis of size frequency distribution of an adult population. Comparison of temperate and tropical records suggests that the duration of the breeding season is regulated by temperature. The minimum age at which larvae achieve competence to settle is approximately the same in temperate and tropical waters, but in both regions development and growth occur at a very variable rate. A number of morphological features which can be considered precursors to settlement are closely associated in their development, which may proceed in response to settlement cues provided by proximity to the substratum. The varying intensity of such cues in different environments may be responsible for delays before some larvae realize competence to settle, accounting for much of the variability in the development process. Accordingly, differences between collections of Indo-West Pacific Lingula larvae can be rationalized; all records are now referred to L. anatina . Extension of pelagic larval life in lingulid larvae appears to involve a strategy different from that of most other marine invertebrates.     

Community regulation: the relative importance of recruitment and predation intensity of an intertidal community dominant in a seascape context

Predicting the strength and context-dependency of species interactions across multiple scales is a core area in ecology. This is especially challenging in the marine environment, where populations of most predators and prey are generally open, because of their pelagic larval phase, and recruitment of both is highly variable. In this study we use a comparative-experimental approach on small and large spatial scales to test the relationship between predation intensity and prey recruitment and their relative importance in shaping populations of a dominant rocky intertidal space occupier, mussels, in the context of seascape (availability of nearby subtidal reef habitat). Predation intensity on transplanted mussels was tested inside and outside cages and recruitment was measured with standard larval settlement collectors. We found that on intertidal rocky benches with contiguous subtidal reefs in New Zealand, mussel larval recruitment is usually low but predation on recruits by subtidal consumers (fish, crabs) is intense during high tide. On nearby intertidal rocky benches with adjacent sandy subtidal habitats, larval recruitment is usually greater but subtidal predators are typically rare and predation is weaker. Multiple regression analysis showed that predation intensity accounts for most of the variability in the abundance of adult mussels compared to recruitment. This seascape-dependent, predation-recruitment relationship could scale up to explain regional community variability. We argue that community ecology models should include seascape context-dependency and its effects on recruitment and species interactions for better predictions of coastal community dynamics and structure.     

Divergent immune priming responses across flour beetle life stages and populations

Growing evidence shows that low doses of pathogens may prime the immune response in many insects, conferring subsequent protection against infection in the same developmental stage (within‐life stage priming), across life stages (ontogenic priming), or to offspring (transgenerational priming). Recent work also suggests that immune priming is a costly response. Thus, depending on host and pathogen ecology and evolutionary history, tradeoffs with other fitness components may constrain the evolution of priming. However, the relative impacts of priming at different life stages and across natural populations remain unknown. We quantified immune priming responses of 10 natural populations of the red flour beetle Tribolium castaneum, primed and infected with the natural insect pathogen Bacillus thuringiensis. We found that priming responses were highly variable both across life stages and populations, ranging from no detectable response to a 13‐fold survival benefit. Comparing across stages, we found that ontogenic immune priming at the larval stage conferred maximum protection against infection. Finally, we found that various forms of priming showed sex‐specific associations that may represent tradeoffs or shared mechanisms. These results indicate the importance of sex‐, life stage‐, and population‐specific selective pressures that can cause substantial divergence in priming responses even within a species. Our work highlights the necessity of further work to understand the mechanistic basis of this variability.     

Depth and timing of settlement of veligers from different populations of giant scallop, Placopecten magellanicus (Gmelin), in thermally stratified mesocosms

The present study was conducted to evaluate the relative roles that water column stratification intensity and possible inter-population behaviour differences play in determining depth of larval settlement of giant scallops, Placopecten magellanicus, in relation to thermoclines. Differences in timing of settlement of larvae from various populations were also examined. Two separate experiments were conducted in a 10.5-m-deep, 3.7-m-diameter, thermally stratified tank with larvae spawned from scallops collected from several adult beds located in areas with differing oceanographic regimes (Georges Bank, Mahone Bay, Passamaquoddy Bay). Previous mesocosm experiments had shown that veligers from these various populations differ in their vertical migration patterns. In the first experiment (December 1992-February 1993), larvae from all three populations were held in separate 9.5-m-deep tubes and exposed to a 1.5 °C temperature differential established over a depth interval of 1 m. The number of settled juveniles (spat) of each population collected at the end of the experiment increased with depth and showed no peak at or above the thermocline. This depth distribution of spat was most likely driven by preferential larval settlement. In the second experiment (February-May 1994), larvae spawned from Georges Bank and Passamaquoddy Bay stocks were held in 9.0-m-deep tubes and exposed to a 5 °C temperature differential established over a depth interval of 1 m. The number of settled spat of both populations was greater above this thermal boundary and increased with decreasing depth. This depth distribution was most likely driven by preferential settlement above the thermocline followed by upward post-settlement migration. The results from the two experiments indicate that larvae from various populations show similar trends in settlement patterns in response to similar thermal stratifications. Stratification intensity, however, does affect depth of larval settlement. In the second experiment, both populations of larvae settled throughout the time interval of collector deployment (larvae 32-82 days old). These results extend the range of planktonic developmental times generally reported in the literature and may be more indicative of natural planktonic development in the field. While Georges Bank larvae settled in consistent numbers through time, Passamaquoddy Bay larvae showed peaks in settlement at certain time periods, indicating that pulses of larval settlement may occur even from an individual spawning event.     

Modelling larval dispersal and behaviour of coral reef fishes

Coral reef fish spend their first few weeks developing in the open ocean, where eggs and larvae appear merciless to tides and currents, before attempting to leave the pelagic zone and settle on a suitable reef. This pelagic dispersal phase is the process that determines population connectivity and allows replenishment of harvested populations across multiple coral reef habitats. Until recently this pelagic larval dispersal phase has been poorly understood and has often been referred to as the ‘black-box’ in the life-history of coral reef fishes. In this perspective article we highlight three areas where mathematical and computational approaches have been used to aid our understanding of this important ecological process. We discuss models that provide insights into the evolution of the pelagic larval phase in coral reef fish, an unresolved question which lends itself well to a modelling approach due to the difficulty in obtaining empirical data on this life history strategy. We describe how studies of fish hearing and physical sound propagation models can be used to predict the detection distance of reefs for settling larval fish, and the potential impact of anthropogenic noise. We explain how random walk models can be used to explore individual- and group-level behaviour in larval fish during the dispersal and settlement stage of their life-history. Finally, we discuss the mutual benefits that mathematical and computational approaches have brought to and gained from the field of larval behaviour and dispersal of reef fishes.     

Habitat structure,conspecific presence and spatial variation in the recruitment of a temperate reef fish

Pronounced spatial variation in recruitment occurs in many marine invertebrate and fish populations and is thought to be critical to the demography of these species. In this study I examined the importance of habitat structure and the presence of conspecific residents to spatial variation in larval settlement and recruitment in a temperate fish Tautogolabrus adspersus. I define settlement as the movement of individuals from the water column to the benthic habitat, while I refer to recruitment as numbers of individuals surviving some arbitrary period of time after settlement. Experiments in which standard habitats were stocked with conspecifics showed that resident conspecifics were not an important factor contributing to small-scale variability in recruitment. Further correlative analyses demonstrated that large-scale variation in recruitment could not be explained by variability in older age classes. By contrast, manipulations of macroalgal structure within a kelp bed demonstrated that recruitment was significantly higher in habitats with a dense understory of foliose and filamentous algae than in habitats with only crustose algae. Understory algae varied in their pattern of disperison among sites, and the dispersion of fish matched that of the plants. In order to determine the effects of differences in patterns of algal dispersion on the demography of associated T. adspersus populations, I used experimental habitat units to manipulate patterns of dispersion. Settlement was significantly greater to randomly placed versus clumped habitats; however, no differences in recruitment between random and clumped habitats were detected. Because recruitment is a function of the numbers of settlers minus the subsequent loss of settlers, rates of mortality or migration must have been higher in the randomly placed habitats. These results are counter to the current paradigm for reef fishes which suggests that larval settlement is the crucial demographic process producing variability in population abundance. In this experiment patterns of settlement were modified by varying the patch structure of the habitat.Contribution number 278 from the Center for Marine Biology, University of New Hampshire     

Larval supply is a good predictor of recruitment in endemic but not non-endemic fish populations at a high latitude coral reef

Despite extensive research, factors influencing the importance of pre- and post-settlement processes to recruitment variability remain ambiguous. Using a novel perspective, we investigated the potential influence of endemism on the relationship between larval supply and recruitment in reef fish populations at Lord Howe Island, Australia. Larval supply and recruitment were measured for three regional endemic and four widespread non-endemic species using light traps, artificial collectors, and underwater visual censuses. Recruitment was correlated with larval supply in endemics but not in non-endemics, likely due to a combination of low larval supply and post-settlement survival of non-endemics. Surveys also indicated that endemics were far more abundant and occurred in more locations than closely related non-endemics. These preliminary findings suggest that either local adaptation enhances recruitment in endemics through higher larval replenishment rates or reduced post-settlement mortality, populations of widespread species at the periphery of their range are poorly adapted to local environmental conditions and therefore experience lower and more variable settlement and post-settlement survival rates, or both.     

Settlement preferences in coral-reef fishes: Effects on patterns of adult and juvenile distributions,individual fitness and population structure

Abstract Coral-reef fishes exhibit a wide range of habitat preferences at settlement. However, the consequences of these preferences to fitness and population dynamics are poorly known. We critically evaluate evidence for these consequences from recent studies of settlement preferences, and note that most studies have concentrated on only a few common taxa. We recommend that this taxonomic base be broadened, and that settlement preferences be incorporated into future studies of the link between larval supply and population dynamics of reef fishes.     

Population Connectivity Shifts at High Frequency within an Open-Coast Marine Protected Area Network

A complete understanding of population connectivity via larval dispersal is of great value to the effective design and management of marine protected areas (MPA). However empirical estimates of larval dispersal distance, self-recruitment, and within season variability of population connectivity patterns and their influence on metapopulation structure remain rare. We used high-resolution otolith microchemistry data from the temperate reef fish Hypsypops rubicundus to explore biweekly, seasonal, and annual connectivity patterns in an open-coast MPA network. The three MPAs, spanning 46 km along the southern California coastline were connected by larval dispersal, but the magnitude and direction of connections reversed between 2008 and 2009. Self-recruitment, i.e. spawning, dispersal, and settlement to the same location, was observed at two locations, one of which is a MPA. Self-recruitment to this MPA ranged from 50–84%; within the entire 60 km study region, self-recruitment accounted for 45% of all individuals settling to study reefs. On biweekly time scales we observed directional variability in alongshore current data and larval dispersal trajectories; if viewed in isolation these data suggest the system behaves as a source-sink metapopulation. However aggregate biweekly data over two years reveal a reef network in which H. rubicundus behaves more like a well-mixed metapopulation. As one of the few empirical studies of population connectivity within a temperate open coast reef network, this work can inform the MPA design process, implementation of ecosystem based management plans, and facilitate conservation decisions.     

Patterns and persistence of larval retention and connectivity in a marine fish metapopulation

Connectivity, the demographic linking of local populations through the dispersal of individuals, is one of the most poorly understood processes in population dynamics, yet has profound implications for conservation and harvest strategies. For marine species with pelagic larvae, direct estimation of connectivity remains logistically challenging and has mostly been limited to single snapshots in time. Here, we document seasonal and interannual patterns of larval dispersal in a metapopulation of the coral reef fish Amphiprion polymnus. A 3‐year record of larval trajectories within and among nine discrete local populations from an area of approximately 35 km was established by determining the natal origin of settled juveniles through DNA parentage analysis. We found that spatial patterns of both self‐recruitment and connectivity were remarkably consistent over time, with a low level of self‐recruitment at the scale of individual sites. Connectivity among sites was common and multidirectional in all years and was not significantly influenced by seasonal variability of predominant surface current directions. However, approximately 75% of the sampled juveniles could not be assigned to parents within the study area, indicating high levels of immigrations from sources outside the study area. The data support predictions that the magnitude and temporal stability of larval connectivity decreases significantly with increasing distance between subpopulations, but increases with the size of subpopulations. Given the considerable effort needed to directly measure larval exchange, the consistent patterns suggest snapshot parentage analyses can provide useful dispersal estimates to inform spatial management decisions.     

Variation in genetic architecture of olfactory behaviour among wild-derived populations of Drosophila melanogaster

Odour-guided behaviour is a quantitative trait determined by many genes that are sensitive to gene-environment interactions. Different natural populations are likely to experience different selection pressures on the genetic underpinnings of chemosensory behaviour. However, few studies have reported comparisons of the quantitative genetic basis of olfactory behaviour in geographically distinct populations. We generated isofemale lines of Drosophila melanogaster from six populations in Argentina and measured larval and adult responses to benzaldehyde. There was significant variation within populations for both larval and adult olfactory behaviour and a significant genotype x sex interaction (GSI) for adult olfactory behaviour. However, there is substantial variation in the contribution of GSI to the total phenotypic variance among populations. Estimates of evolvability are orders of magnitude higher for larvae than for adults. Our results suggest that the potential for evolutionary adaptation to the chemosensory environment is greater at the larval feeding stage than at the adult reproductive stage.     

Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Intra-annual genetic variation in the downstream larval drift of sutchi catfish (Pangasianodon hypophthalmus) in the Mekong river

Larvae of the sutchi catfish Pangasianodon hypophthalmus were collected during peak downstream drift in the Lower Mekong river on four occasions over an 8-week period during the 2003 spawning season, and genotyped using seven microsatellite loci. We provide evidence for several heterogeneous groups within and among the temporally discrete larval peak samples. Strong evidence for a significant deficit of heterozygotes was observed for each larval sample and the pooled sample, possibly due to population admixture. Although individual-based assignment tests suggested that each larval peak sample was admixed, significant but low genetic differentiation was observed among larval samples ( F _ST = 0.0052, P  < 0.01). The lack of significant relatedness confirms the multifamily composition of each larval group, excluding family bias to explain the observed genetic heterogeneity. Both the entire larval peak and each temporally separated larval peak originated from spawning groups with heterogeneous allelic composition involving several distinct spawning events. We propose three explanations to account for our findings: (1) the ecological match/mismatch hypothesis; (2) the genetic 'sweepstakes' selection hypothesis; and (3) life-history-specific characteristics of the spawning populations. Finally, an intra-annual shift in the contribution of the spawning populations to the larval drift was detected on successive occasions.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 719–728.     

The significance of variable and density-independent post-recruitment mortality in local populations of reef fishes

Abstract In this paper I focus on how post-settlement mortality may modify initial patterns of settlement in reef fish. Infrequent recruitment surveys may underestimate the role of early post-settlement mortality as most mortality in reef fishes occurs shortly after settlement. Consequently, results from infrequent recruitment surveys shed little light on the mechanisms producing patterns of abundance because these surveys ignore early post-settlement mortality. Variation in density-independent mortality may be a common mechanism that can prevent a positive relationship between larval settlement and subsequent population abundance. Although density-dependent mortality is the most commonly recognized mechanism that can disrupt the correlation between settlement and adult abundance, density-independent mortality’ can also destroy this correlation if the variance associated with post-settlement mortality is greater than variance in settlement. This point is illustrated with a simulation model in which I modelled two populations: a piscivorous fish population that was recruitment-limited with constant mortality, and a prey population that had variable recruitment and mortality that was a function of the size of the predator population. The results of this model indicate that even when mortality of prey is density-independent, predation can determine prey abundance when variation in piscivore recruitment is high relative to prey recruitment. Thus, initial patterns of prey settlement can be modified by a recruitment-limited predator population.     

A model study on variations in larval supply: are populations of the polychaete <Emphasis Type="Italic">Owenia fusiformis</Emphasis> in the English Channel open or closed?

The polychaete Owenia fusiformis is one of the most ecologically important species in the muddy fine sand sediments in the English Channel where it is distributed in geographically separated populations. A vertically averaged Lagrangian hydrodynamic model integrating tidal residual currents and wind-induced currents was used to drive an advection–diffusion model for investigating the variability of larval transport in order to assess the self-seeding capabilities and the degree of connectivity between local populations. Three different types of environmental forcing (i.e. tidal forcing alone, tidal forcing coupled with either NE winds or SW winds) were applied to 19 distinct populations. Without wind influence, self-seeding is the principal mechanism involved in the renewal of most populations. However, larval retention ranged from under 1% up to 81% in relation to the adult habitat size and the mean velocity of tidal residual currents. Wind forcing had a strong influence on larval dispersal patterns by modifying the origin and densities of settlers as well as the degree of connectivity between populations. As a consequence, larval supply from distant populations generally exceeded local supply and the inter-annual variability of wind forcing induced large year-to-year variations in larval settlement rates. Larval exchanges occurred mainly between neighbouring populations and three groups of interconnected local populations were thereby identified. Within each group, settlement patterns were related to inter-annual variations in the direction and magnitude of larval exchanges. Electronic Publication