Could Seals Prevent Cod Recovery in the Baltic Sea?

Fish populations are increasingly affected by multiple human and natural impacts including exploitation, eutrophication, habitat alteration and climate change. As a result many collapsed populations may have to recover in ecosystems whose structure and functioning differ from those in which they were formerly productive and supported sustainable fisheries. Here we investigate how a cod (Gadus morhua) population in the Baltic Sea whose biomass was reduced due to a combination of high exploitation and deteriorating environmental conditions might recover and develop in the 21st century in an ecosystem that likely will change due to both the already started recovery of a cod predator, the grey seal Halichoerus grypus, and projected climate impacts. Simulation modelling, assuming increased seal predation, fishing levels consistent with management plan targets and stable salinity, shows that the cod population could reach high levels well above the long-term average. Scenarios with similar seal and fishing levels but with 15% lower salinity suggest that the Baltic will still be able to support a cod population which can sustain a fishery, but biomass and yields will be lower. At present knowledge of cod and seal interactions, seal predation was found to have much lower impact on cod recovery, compared to the effects of exploitation and salinity. These results suggest that dual management objectives (recovery of both seal and cod populations) are realistic but success in achieving these goals will also depend on how climate change affects cod recruitment.     

Connectivity,persistence, and loss of high abundance areas of a recovering marine fish population in the Northwest Atlantic Ocean

In the early 1990s, the Northwest Atlantic Ocean underwent a fisheries‐driven ecosystem shift. Today, the iconic cod (Gadus morhua) remains at low levels, while Atlantic halibut (Hippoglossus hippoglossus) has been increasing since the mid‐2000s, concomitant with increasing interest from the fishing industry. Currently, our knowledge about halibut ecology is limited, and the lack of recovery in other collapsed groundfish populations has highlighted the danger of overfishing local concentrations. Here, we apply a Bayesian hierarchical spatiotemporal approach to model the spatial structure of juvenile Atlantic halibut over 36 years and three fisheries management regimes using three model parameters to characterize the resulting spatiotemporal abundance structure: persistence (similarity of spatial structure over time), connectivity (coherence of temporal pattern over space), and spatial variance (variation across the seascape). Two areas of high juvenile abundance persisted through three decades whereas two in the northeast are now diminished, despite the increased abundance and landings throughout the management units. The persistent areas overlap with full and seasonal area closures, which may act as refuges from fishing. Connectivity was estimated to be 250 km, an order of magnitude less than the distance assumed by the definition of the Canadian management units (~2,000 km). The underlying question of whether there are distinct populations within the southern stock unit cannot be answered with this model, but the smaller ~250 km scale of coherent temporal patterns suggests more complex population structure than previously thought, which should be taken into consideration by fishery management.     

Evolutionary response to size-selective mortality in an exploited fish population

Many collapsed fish populations have failed to recover after a decade or more with little fishing. This may reflect evolutionary change in response to the highly selective mortality imposed by fisheries. Recent experimental work has demonstrated a rapid genetic change in growth rate in response to size-selective harvesting of laboratory fish populations. Here, we use a 30-year time-series of back-calculated lengths-at-age to test for a genetic response to size-selective mortality in the wild in a heavily exploited population of Atlantic cod (Gadus morhua). Controlling for the effects of density- and temperature-dependent growth, the change in mean length of 4-year-old cod between offspring and their parental cohorts was positively correlated with the estimated selection differential experienced by the parental cohorts between this age and spawning. This result supports the hypothesis that there have been genetic changes in growth in this population in response to size-selective fishing. Such changes may account for the continued small size-at-age in this population despite good conditions for growth and little fishing for over a decade. This study highlights the need for management regimes that take into account the evolutionary consequences of fishing.     

Demographic analyses of the dusky shark, <Emphasis Type="Italic">Carcharhinus obscurus,</Emphasis> in the Northwest Atlantic incorporating hooking mortality estimates and revised reproductive parameters

Demographic analyses were conducted for the dusky shark, Carcharhinus obscurus, in the Northwest Atlantic. Fishery-independent, fishery-dependent, and previously published data were used to estimate age specific by-catch mortality and reproductive cycle for incorporation into the analyses. Correspondence analyses were conducted for viability (dead or alive), shark length, month, soaktime, and region. Dead sharks were associated with small size, long soak times, and the spring season, while live sharks were associated with large size and shorter soak times. Data on reproduction suggested a 3-year reproductive cycle consisting of a 2-year gestation period and a 1-year resting period. Litter sizes ranged from 3–12 embryos. Decreasing age-specific fishing mortality and a 3-year reproductive cycle were used in age structured life tables which incorporated stochasticity in life history parameters through Monte Carlo simulation to estimate annual population growth. Demographic analyses showed that the dusky shark population would decline even at low levels of fishing mortality despite low natural mortality for neonates. The demographics of this species make it among the most vulnerable to excessive fishing mortality, and suggest stringent regulatory measures may be required to recover collapsed populations.     

A century of fish growth in relation to climate change,population dynamics and exploitation

Marine ecosystems, particularly in high‐latitude regions such as the Arctic, have been significantly affected by human activities and contributions to climate change. Evaluating how fish populations responded to past changes in their environment is helpful for evaluating their future patterns, but is often hindered by the lack of long‐term biological data available. Using otolith increments of Northeast Arctic cod (Gadus morhua) as a proxy for individual growth, we developed a century‐scale biochronology (1924–2014) based on the measurements of 3,894 fish, which revealed significant variations in cod growth over the last 91 years. We combined mixed‐effect modeling and path analysis to relate these growth variations to selected climate, population and fishing‐related factors. Cod growth was negatively related to cod population size and positively related to capelin population size, one of the most important prey items. This suggests that density‐dependent effects are the main source of growth variability due to competition for resources and cannibalism. Growth was also positively correlated with warming sea temperatures but negatively correlated with the Atlantic Multidecadal Oscillation, suggesting contrasting effects of climate warming at different spatial scales. Fishing pressure had a significant but weak negative direct impact on growth. Additionally, path analysis revealed that the selected growth factors were interrelated. Capelin biomass was positively related to sea temperature and negatively influenced by herring biomass, while cod biomass was mainly driven by fishing mortality. Together, these results give a better understanding of how multiple interacting factors have shaped cod growth throughout a century, both directly and indirectly.     

Characterization of new SSR‐EST markers in cod,Gadus morhua

For years, fishermen have noted that cod (Gadus morhua) within the Georges Bank Atlantic fishery spawn multiple times annually with peaks in the spring and in the fall. To evaluate the hypothesis that these differences in reproductive behaviour reflect genetically different spawning populations, we identified eight polymorphic simple sequence repeats within expressed sequence tags (ESTs). Previously, only a few genetic markers have been informative for distinguishing between cod populations of the Northwest Atlantic. The development of additional markers from ESTs will be a valuable tool to improve our understanding of population structure in this commercially important species.     

Maladaptive changes in multiple traits caused by fishing: impediments to population recovery

Some overharvested fish populations fail to recover even after considerable reductions in fishing pressure. The reasons are unclear but may involve genetic changes in life history traits that are detrimental to population growth when natural environmental factors prevail. We empirically modelled this process by subjecting populations of a harvested marine fish, the Atlantic silverside, to experimental size-biased fishing regimes over five generations and then measured correlated responses across multiple traits. Populations where large fish were selectively harvested (as in most fisheries) displayed substantial declines in fecundity, egg volume, larval size at hatch, larval viability, larval growth rates, food consumption rate and conversion efficiency, vertebral number, and willingness to forage. These genetically based changes in numerous traits generally reduce the capacity for population recovery.     

Phylogeographic mitogenomics of Atlantic cod Gadus morhua: Variation in and among trans‐Atlantic,trans‐Laurentian,Northern cod,and landlocked fjord populations

The historical phylogeography, biogeography, and ecology of Atlantic cod (Gadus morhua) have been impacted by cyclic Pleistocene glaciations, where drops in sea temperatures led to sequestering of water in ice sheets, emergence of continental shelves, and changes to ocean currents. High‐resolution, whole‐genome mitogenomic phylogeography can help to elucidate this history. We identified eight major haplogroups among 153 fish from 14 populations by Bayesian, parsimony, and distance methods, including one that extends the species coalescent back to ca. 330 kya. Fish from the Barents and Baltic Seas tend to occur in basal haplogroups versus more recent distribution of fish in the Northwest Atlantic. There was significant differentiation in the majority of trans‐Atlantic comparisons (Φ_ST = .029–.180), but little or none in pairwise comparisons within the Northwest Atlantic of individual populations (Φ_ST = .000–.060) or defined management stocks (Φ_ST = .000–.023). Monte Carlo randomization tests of population phylogeography showed significantly nonrandom trans‐Atlantic phylogeography versus absence of such structure within various partitions of trans‐Laurentian, Northern cod (NAFO 2J3KL) and other management stocks, and Flemish Cap populations. A landlocked meromictic fjord on Baffin Island comprised multiple identical or near‐identical mitogenomes in two major polyphyletic clades, and was significantly differentiated from all other populations (Φ_ST = .153–.340). The phylogeography supports a hypothesis of an eastern origin of genetic diversity ca. 200–250 kya, rapid expansion of a western superhaplogroup comprising four haplogroups ca. 150 kya, and recent postglacial founder populations.     

It is the economy,stupid! Projecting the fate of fish populations using ecological–economic modeling

Four marine fish species are among the most important on the world market: cod, salmon, tuna, and sea bass. While the supply of North American and European markets for two of these species – Atlantic salmon and European sea bass – mainly comes from fish farming, Atlantic cod and tunas are mainly caught from wild stocks. We address the question what will be the status of these wild stocks in the midterm future, in the year 2048, to be specific. Whereas the effects of climate change and ecological driving forces on fish stocks have already gained much attention, our prime interest is in studying the effects of changing economic drivers, as well as the impact of variable management effectiveness. Using a process‐based ecological–economic multispecies optimization model, we assess the future stock status under different scenarios of change. We simulate (i) technological progress in fishing, (ii) increasing demand for fish, and (iii) increasing supply of farmed fish, as well as the interplay of these driving forces under different scenarios of (limited) fishery management effectiveness. We find that economic change has a substantial effect on fish populations. Increasing aquaculture production can dampen the fishing pressure on wild stocks, but this effect is likely to be overwhelmed by increasing demand and technological progress, both increasing fishing pressure. The only solution to avoid collapse of the majority of stocks is institutional change to improve management effectiveness significantly above the current state. We conclude that full recognition of economic drivers of change will be needed to successfully develop an integrated ecosystem management and to sustain the wild fish stocks until 2048 and beyond.     

Fishing,reproductive volume and regulation: population dynamics and exploitation of the eastern Baltic cod

The relative importance of exploitation rate and environmental variability in generating fluctuations of harvested populations is a key issue in academic ecology as well as population management. We studied how the eastern Baltic cod (Gadus morhua) is affected by fishing and environmental variation by using a newly developed single species state-space model. Survey data and auxiliary environmental data were used to estimate the model parameters. The model was then used to predict future development of the eastern Baltic cod under different fishing mortalities and abiotic conditions. Abiotic condition was represented by an index: reproductive volume which is the volume of water suitable (in terms of salinity and oxygen content) for the successful development of the early life stages of Baltic cod. The model included direct density dependence, fishing, and a lagged effect of reproductive volume. Our analysis showed that fishing rate is approximately three times more important than reproductive volume in explaining the population dynamics. Furthermore, our model suggests either under- or over-compensatory dynamics depending on the reproductive volume and long term catch levels. It follows that fishing can either reduce or increase temporal oscillations of the cod stock depending on whether the dynamics is over- or undercompensatory, respectively. The sustainable level of fishing rate is however dependent on reproductive volume. Our model predicts a dual role of fishing rate, stabilizing when reproductive volume is high and destabilizing when it is low. Exploitation rate may therefore increase or decrease the risk of the population of cod dropping below a given biomass reference point depending on the environmental conditions, which has practical implications for fisheries management.     

Long-term studies on genetic interaction between wild and ranched cod Gadus morhua by use of a genetic marked strain

Releases of farmed fish, whether accidental from commercial aquaculture facilities or intentional as part of stock enhancement/ranching activities, are considered to pose a risk to native gene pools. Stock enhancement studies of Atlantic cod, Gadus morhua , based on artificially produced juveniles, were initiated in 1984 in western Norway, and genetic aspects were incorporated. In order to investigate potential interbreeding between released and wild cod, a genetically marked cod strain was developed, being homozygotic for a rare allele ( GPI‐1*30 ) expressed in white muscle tissue. In the period from 1990 to 1994, juveniles from the genetic marked strain were released in large quantities in three locations (Masfjord, Øygarden, Heimarkspollen), giving a significant increase of the marker allele in the local wild cod populations. Recently, studies have been conducted in the same areas to estimate the extent of interbreeding between the wild and released cod. The results, however, revealed no permanent increase of the frequency of the marker allele and/or GPI‐1*30 heterozygotes as would be expected from interbreeding. The recent data are compared with comprehensive genetic data of the cod populations in the areas before the actual releases, covering the full period from 1994 to 2003. The present results are also discussed in relation to fishing pressure on coastal cod, migration information and reproductive success of released, genetically marked cod.     

Transient demographic dynamics of recovering fish populations shaped by past climate variability,harvest, and management

Large-scale commercial harvesting and climate-induced fluctuations in ocean properties shape the dynamics of marine populations as interdependent drivers at varied timescales. Persistent selective removals of larger, older members of a population can distort its demographic structure, eroding resilience to fluctuations in habitat conditions and thus amplifying volatility in transient dynamics. Many historically depleted marine fish stocks have begun showing signs of recovery in recent decades following the implementation of stricter management measures. But these interventions coincide with accelerated changes in the oceans triggered by increasingly warmer, more variable climates. Applying multilevel models to annual estimates of demographic metrics of 38 stocks comprising 11 species across seven northeast Atlantic ecoregions, this study explores how time-varying local and regional climates contributed to the transient dynamics of recovering populations exposed to variable fishing pressures moderated by management actions. Analyses reveal that progressive reductions in fishing pressure and shifting climate conditions discontinuously shaped rebuilding patterns of the stocks through restorations of maternal demographic structure (reversing age truncation) and reproductive capacity. As the survival rate and demographic structure of reproductive fish improved, transient growth became less sensitive to variability in recruitment and juvenile survival and more to that in adult survival. As the biomass of reproductive fish rose, recruitment success also became increasingly regulated by density-dependent processes involving higher numbers of older fish. When reductions in fishing pressure were insufficient or delayed, however, stocks became further depleted, with more eroded demographic structures. Although warmer local climates in spawning seasons promoted recruitment success in some ecoregions, changing climates in recent decades began adversely affecting reproductive performances overall, amplifying sensitivities to recruitment variability. These shared patterns underscore the value of demographic transients in developing robust strategies for managing marine resources. Such strategies could form the foundation for effective applications of adaptive measures resilient to future environmental change.     

Parrotfish sex ratios recover rapidly in Bermuda following a fishing ban

Parrotfishes are an ecologically and commercially important teleost group whose grazing contributes to maintaining coral-dominated states on hermatypic reefs. However, overfishing has skewed sex ratios of Atlantic parrotfishes because fishing has disproportionate impacts on larger individuals, and males are generally larger than females. Whether protection from fishing may allow sex ratios to return to equilibrium is unknown, as fishing can induce irreversible ecological and/or evolutionary shifts. Bermuda banned trap fishing in 1990, creating a unique opportunity to analyse long-term responses of Atlantic parrotfishes to release from fishing. We found that sex ratios of four common parrotfishes were initially skewed, with male proportions ranging from 0.04 to 0.18. However, male proportions rebounded within 3–4 yr, equilibrating at values ranging from 0.36 to 0.54, similar to those reported at unfished sites in the region. Our results are encouraging for regional efforts to recover lost grazing function by restoring overfished herbivore populations.     

Fishing,predation, and temperature drive herring decline in a large marine ecosystem

Since 1960, landings of Atlantic herring have been the greatest of any marine species in Canada, surpassing Atlantic cod and accounting for 24% of the total seafood harvested in Atlantic Canada. The Scotian Shelf‐Bay of Fundy herring fisheries (NAFO Division 4VWX) is among Canada''s oldest and drives this productivity, accounting for up to 75% of the total herring catch in some years. The stocks’ productivity and overall health have declined since 1965. Despite management measures to promote recovery implemented since 2003, biomass remains low and is declining. The factors that drive the productivity of 4VWX herring are primarily unresolved, likely impeding the effectiveness of management actions on this stock. We evaluated potential drivers of herring variability by analyzing 52 time‐series that describe the temporal and spatial evolution of the 4VWX herring population and the physical, ecological, and anthropogenic factors that could affect them using structural equation models. Variation in herring biomass was best accounted for by the exploitation rate''s negative effect and the geographic distribution of fishing and recruitment. Thermal phenology and temperature adversely and egg predation positively impacted the early life stage mortality rate and, ultimately, adult biomass. These findings are broadly relevant to fisheries management, but particularly for 4VWX herring, where the current management approach does not consider their early life stage dynamics or assess them within the ecosystem or climate change contexts.     

Consequences of fisheries-induced evolution for population productivity and recovery potential

Fisheries-induced evolution has become a major branch of the research on anthropogenic and contemporary evolution. Within the conservation context, fisheries-induced evolution has been hypothesized to negatively affect the persistence and recovery potential of depleted populations, but this has not been explicitly investigated. Here, we investigate how fisheries-induced evolution of Atlantic cod (Gadus morhua L.) life histories affects per capita population growth rate, a parameter negatively correlated with extinction risk. We simulate the evolutionary and ecological dynamics of a cod population for a 100 year period of size-selective harvesting, followed thereafter by 300 years of recovery. To evaluate the relative importance of harvest-induced evolution, we either allowed life histories to evolve during and after the fishing period, or we assumed that fisheries-induced evolution was absent. Population growth rates did not differ appreciably between the evolutionary and non-evolutionary simulation scenarios, despite the emergence of rather pronounced differences in life histories. The underlying reason was that in the absence of fishing the cumulative lifetime reproductive outputs were very similar among differing life histories. The results suggest that fisheries-induced evolution might not always have as clear-cut an effect on population growth rate as previously anticipated.     

Long‐term change in a behavioural trait: truncated spawning distribution and demography in Northeast Arctic cod

Harvesting may be a potent driver of demographic change and contemporary evolution, which both may have great impacts on animal populations. Research has focused on changes in phenotypic traits that are easily quantifiable and for which time series exist, such as size, age, sex, or gonad size, whereas potential changes in behavioural traits have been under‐studied. Here, we analyse potential drivers of long‐term changes in a behavioural trait for the Northeast Arctic stock of Atlantic cod Gadus morhua, namely choice of spawning location. For 104 years (1866–1969), commercial catches were recorded annually and reported by county along the Norwegian coast. During this time period, spawning ground distribution has fluctuated with a trend towards more northerly spawning. Spawning location is analysed against a suite of explanatory factors including climate, fishing pressure, density dependence, and demography. We find that demography (age or age at maturation) had the highest explanatory power for variation in spawning location, while climate had a limited effect below statistical significance. As to potential mechanisms, some effects of climate may act through demography, and explanatory variables for demography may also have absorbed direct evolutionary change in migration distance for which proxies were unavailable. Despite these caveats, we argue that fishing mortality, either through demographic or evolutionary change, has served as an effective driver for changing spawning locations in cod, and that additional explanatory factors related to climate add no significant information.     

Mixed stock analysis and the power of different classes of molecular markers in discriminating coastal and oceanic Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) on the Lofoten spawning grounds,Northern Norway

Atlantic cod (Gadus morhua) encompasses many different populations or stocks, which in part are managed separately. In the northeast Atlantic cod is divided into two main management units; northeast Arctic cod and coastal cod. These two groups have traditionally been separated by otolith classification. In this study, the power of different classes of genetic markers in separating the two cod groups was investigated. The variation in thirteen genetic markers, including allozymes, haemoglobin, the scDNA locus Pantophysin (Pan I) and a number of microsatellites was investigated, and mixed stock analysis and individual assignment tests were performed on samples comprising a mixture of individuals of putative coastal and oceanic type cod. The genetic analyses showed a large genetic differentiation between outer stations and stations located closer to the mainland shore. Mixed stock analysis and individual assignment tests used for estimation of stock proportions gave results similar to those obtained by otolith classification. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

The rise of a marine generalist predator and the fall of beta diversity

Determining the importance of physical and biological drivers in shaping biodiversity in diverse ecosystems remains a global challenge. Advancements have been made towards this end in large marine ecosystems with several studies suggesting environmental forcing as the primary driver. However, both empirical and theoretical studies point to additional drivers of changes in diversity involving trophic interactions and, in particular, predation. Moreover, a more integrated but less common approach to the assessment of biodiversity changes involves analyses of spatial β diversity, whereas most studies to date assess only changes in species richness (α diversity). Recent research has established that when cod, a dominant generalist predator, was overfished and collapsed in a northwest Atlantic food web, spatial β diversity increased; that is, the spatial structure of the fish assemblage became increasingly heterogeneous. If cod were to recover, would this situation be reversible, given the inherent complexity and non‐linear dynamics that typify such systems? A dramatic increase of cod in an ecologically similar large marine ecosystem may provide an answer. Here we show that spatial β diversity of fish assemblages in the Barents Sea decreased with increasing cod abundance, while decadal scale changes in temperature did not play a significant role. These findings indicate a reversibility of the fish assemblage structure in response to changing levels of an apex predator and highlight the frequently overlooked importance of trophic interactions in determining large‐scale biodiversity patterns. As increased cod abundance was largely driven by changes in fisheries management, our study also shows that management policies and practices, particularly those involving apex predators, can have a strong effect in shaping spatial diversity patterns, and one should not restrict the focus to effects of climate change alone.     

Shark bycatch and depredation in the U.S. Atlantic pelagic longline fishery

The non-target bycatch of sharks in pelagic longline (PLL) fisheries represents a potential source of compromise to shark populations worldwide. Moreover, shark bycatch and depredation (damage inflicted on gear, bait, and catch) complicates management of sharks and other species, and can undermine the operations and financial interests of the pelagic longline industry. Thus, deducing means to reduce shark interactions is in the best interest of multiple stakeholder groups. Prior to doing so, however, the extent, cause and effect of these interactions must be better understood. In this review we address or conduct the following in relation to the U.S. Atlantic, Gulf of Mexico and Caribbean PLL fishery: (1) U.S. management governing shark interactions in the Atlantic; (2) the primary species encountered and historical shark catch data associated with PLL fishing in the Atlantic; (3) a historical comparison of area-specific shark species catch records between the two primary sources of shark catch data in this fishery; (4) the conditions and dynamics that dictate shark interactions in this fishery, and potential means to reduce these interactions, and; (5) a synthesis of the estimated impacts of this fishery on shark populations relative to other fisheries in the Atlantic. As has been found in other PLL fisheries, the blue shark (Prionace glauca) is clearly the shark species most commonly encountered in this fishery in the Atlantic, and receives the majority of attention in this review. U.S. management areas with high relative shark species diversities had a greater divergence in historical shark species percent-compositions between data sources (Pelagic Observer Program versus mandatory pelagic Logbook databases); this complicates the ability to conclude which species are most impacted by PLL fishing in those areas. The current fishing effort by the U.S. PLL fleet is small compared to that of PLL fishing targeting sharks in the Atlantic by non-U.S. fleets, and therefore poses a comparatively lower threat to the stability of Atlantic shark populations. However, incidental shark encounters are inevitable in U.S. Atlantic PLL fishing operations. Thus, it is in the best interest of all stakeholders in the Atlantic to better understand the extent and conditions governing these interactions, and to explore methods to reduce both their occurrence and those aspects leading to higher rates of incidental shark mortality.     

Cod Gadus morhua L. populations as behavioural units: inference from time series on juvenile abundance in the eastern Skagerrak

Abundance and distribution of cod Gadus morhua in various size intervals and age groups between 2000 and 2005 were followed in coastal trawl surveys. In spite of a reduction in fishing pressure in recent years and high cod recruitment in the Skagerrak region in 2001 and 2003, no recovery could be evidenced. The survey data clearly showed that low cod density areas were not recolonized, even though abundance of juvenile cod remained high for about a year after the recruitment episodes. Increased abundance of fish >400 mm total length was only discernible at some scattered locations where other studies also have suggested local populations still to be present. The intermittent high recruitment has been linked to an inflow of egg and larvae from the North Sea, a theory which also has gained support from genetic studies. It was thus argued that the disappearance of the juvenile cod from the inshore is an effect of a migratory behaviour; the fish of offshore origin eventually leave the coast for the open Skagerrak or the North Sea. These findings support a view on cod populations as essentially behavioural entities, whereas dispersal of early life stages may be less important as a structuring mechanism.