Importance du sédiment dans la distribution des Lingules

Log-probability curves of grain-size distribution have been analysed of sediments collected in three previously described localities occupied by either Lingula anatina or L. reevei (Brachiopoda, Inarticulata). The grain-size population transported by saltation (average about 92–223 μm), generally associated with traction load population, determines Lingula distribution. The two preferential substrates of both studied species (density > 100 individuals/m^-2) are compact and stable sediments under moderate water currents: either coarse sands and gravels clogged by fine and very fine sands, or fine, very fine and clay sands (saltation populations more than 60%). As soon as the suspension population and/or traction increase in the sediment to the detriment of the saltation population, the density of Lingula decreases rapidly. Nevertheless, if grain-size distribution and populations have a primordial function for Lingula , other ecological features (i.e. the environmental fauna, high occurrence of digging species, the available nutrients) will affect the distribution and sometimes become prevailing. The absence of important changes in the shell shape of Lingula since its origin suggests that the grain-size requirements remained about the same.     

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.     

Population structure, growth and distribution of Lingula anatina (Brachiopoda) in Queensland, Australia

Size-frequency data collected in large samples from five populations of Lingula anatina from North Queensland intertidal sandflats conformed to previously known brachiopod size-frequency distributions. Probability paper analysis of the samples and constituent sub-samples confirmed that all the distributions had a hidden poly-modality resulting from a long but defined breeding season. Two patterns of annual recruitment were revealed; the first is the periodic failure of the populations to gain recruits and the second is one of patchy distribution of settlement cohorts. These patterns introduce sampling bias into the derivation of lingulid size frequency curves unless compensated by more extensive sampling methods than have previously been considered adequate. The interpretation of year classes has enabled a growth curve to be derived. The populations inhabited predominantly sandy sub-strates and displayed aggregated distributions. Littoral distribution of the species is regarded as optimal rather than marginal.     

Relative impacts of adult movement, larval dispersal and harvester movement on the effectiveness of reserve networks

Movement of individuals is a critical factor determining the effectiveness of reserve networks. Marine reserves have historically been used for the management of species that are sedentary as adults, and, therefore, larval dispersal has been a major focus of marine-reserve research. The push to use marine reserves for managing pelagic and demersal species poses significant questions regarding their utility for highly-mobile species. Here, a simple conceptual metapopulation model is developed to provide a rigorous comparison of the functioning of reserve networks for populations with different admixtures of larval dispersal and adult movement in a home range. We find that adult movement produces significantly lower persistence than larval dispersal, all other factors being equal. Furthermore, redistribution of harvest effort previously in reserves to remaining fished areas ('fishery squeeze') and fishing along reserve borders ('fishing-the-line') considerably reduce persistence and harvests for populations mobile as adults, while they only marginally changes results for populations with dispersing larvae. Our results also indicate that adult home-range movement and larval dispersal are not simply additive processes, but rather that populations possessing both modes of movement have lower persistence than equivalent populations having the same amount of 'total movement' (sum of larval and adult movement spatial scales) in either larval dispersal or adult movement alone.     

Larval dispersal dampens population fluctuation and shapes the interspecific spatial distribution patterns of rocky intertidal gastropods

Many marine benthic invertebrates pass through a planktonic larval stage whereas others spend their entire lifetimes in benthic habitats. Recent studies indicate that non‐planktonic species show relatively greater fine‐scale patchiness than do planktonic species, but the underlying mechanisms remain unknown. One hypothesis for such a difference is that larval dispersal enhances the connectivity of populations and buffers population fluctuations and reduces local extinction risk, consequently increasing patch occupancy rate and decreasing spatial patchiness. If this mechanism does indeed play a significant role, then the distribution of non‐planktonic species should be more aggregated – both temporally and spatially – than the distribution of species with a planktonic larval stage. To test this prediction, we compared 1) both the spatial and the temporal abundance–occupancy relationships and 2) both the spatial and the temporal mean–variance relationships of population size across species of rocky intertidal gastropods with differing dispersive traits from the Pacific coast of Japan. We found that, compared to planktonic species, non‐planktonic species exhibited 1) a smaller occupancy rate for any given level of mean population size and 2) greater variations in population size, both spatially and temporally. This suggests that the macroecological patterns observed in this study (i.e. the abundance–occupancy relationships and mean–variance relationships of population size across species) were shaped by the effect of larval dispersal dampening population fluctuation, which works over both space and time. While it has been widely assumed that larval dispersal enhances population fluctuations, larval dispersal may in fact enhance the connectively of populations and buffer population fluctuations and reduce local extinction risks.     

Genetic population structure of the endemic fourline wrasse (Larabicus quadrilineatus) suggests limited larval dispersal distances in the Red Sea

The connectivity among marine populations is determined by the dispersal capabilities of adults as well as their eggs and larvae. Dispersal distances and directions have a profound effect on gene flow and genetic differentiation within species. Genetic homogeneity over large areas is a common feature of coral reef fishes and can reflect high dispersal capability resulting in high levels of gene flow. If fish larvae return to their parental reef, gene flow would be restricted and genetic differentiation could occur. Larabicus quadrilineatus (Labridae) is considered as an endemic fish species of the Red Sea and Gulf of Aden. The juveniles of this species are cleaner fish that feed on ectoparasites of other fishes. Here, we investigated the genetic population structure and gene flow in L. quadrilineatus among five locations in the Red Sea to infer connectivity among them. To estimate genetic diversity, we analysed 369 bp of 237 mitochondrial DNA control region sequences. Haplotype and nucleotide diversities were higher in the southern than in the northern Red Sea. Analysis of molecular variance (amova) detected the highest significant genetic variation between northern and central/southern populations (Phi(CT) = 0.01; P < 0.001). Migration analysis revealed a several fold higher northward than southward migration, which could be explained by oceanographic conditions and spawning season. Even though the Phi(ST) value of 0.01 is rather low and implies a long larval dispersal distance, estimates based on the isolation-by-distance model show a very low mean larval dispersal distance (0.44-5.1 km) compared to other studies. In order to enable a sustainable ornamental fishery on the fourline wrasse, the results of this study suggest that populations in the northern and southern Red Sea should be managed separately as two different stocks. The rather low larval dispersal distance of about 5 km needs to be considered in the design of marine protected areas to enable connectivity and self-seeding.     

The recovery of coral genetic diversity in the Sunda Strait following the 1883 eruption of Krakatau

Surveys of microsatellite variation show that genetic diversity has largely recovered in two reef-building corals, Pocillopora damicornis and Seriatopora hystrix (Scleractinia: Pocilloporidae), on reefs which were decimated by the eruption of the volcano Krakatau in 1883. Assignment methods and gene flow estimates indicate that the recolonization of Krakatau occurred mainly from the closest upstream reef system, Pulau Seribu, but that larval input from other regions has also occurred. This pattern is clearer in S. hystrix, which is traditionally the more dispersal-limited species. Despite these observed patterns of larval dispersal, self-recruitment appears to now be the most important factor in supplying larvae to coral populations in Krakatau. This suggests that the colonization of devastated reefs can occur quickly through larval dispersal; however, their survival requires local sources of larvae for self-recruitment. This research supports the observation that the recovery of genetic diversity in coral reef animals can occur on the order of decades and centuries rather than millennia. Conservation measures aimed at sustaining coral reef populations in Krakatau and elsewhere should include both the protection of upstream source populations for larval replenishment should disaster occur as well as the protection of large adult colonies to serve as local larval sources.     

Fine scale dispersal in Banggai Cardinalfish, Pterapogon kauderni, a coral reef species lacking a pelagic larval phase

Dispersal in marine species results from complex interactions between biotic and abiotic factors. Importantly, the pelagic larval phase of most marine species adds a significant degree of complexity. Therefore, a growing body of work is focusing on those rare species that lack a pelagic larval phase (usually brooding species). For such species, large-scale gene flow has been shown to be very low, thus following the expectation of a relationship between realized dispersal and pelagic larval duration. Yet, little is known about the dispersal of those species at very small geographic scales. In this study, we focused on the Banggai Cardinalfish, Pterapogon kauderni, a mouthbrooding species that lacks a pelagic larval phase. Based on previously identified microsatellites, we scored 12 populations around the southern island of Bangkulu, in the Banggai Archipelago, Indonesia. While only 60 km in perimeter, we found that this island harbors very distinct populations of P. kauderni. Indeed, assignment tests self-assigned 10 out of those 12 populations. These results mirror the very high level of self-assignment at the level of the entire archipelago, where, out of 13 populations, 70% of the individuals were reassigned to their source population. Therefore, our data show consistency between small and large-scale dispersal. In addition, in light of the recent expansion in the harvesting of this species for the pet trade, our data have important conservation implications.     

A multispecies approach reveals hot spots and cold spots of diversity and connectivity in invertebrate species with contrasting dispersal modes

Genetic diversity is crucial for species’ maintenance and persistence, yet is often overlooked in conservation studies. Species diversity is more often reported due to practical constraints, but it is unknown if these measures of diversity are correlated. In marine invertebrates, adults are often sessile or sedentary and populations exchange genes via dispersal of gametes and larvae. Species with a larval period are expected to have more connected populations than those without larval dispersal. We assessed the relationship between measures of species and genetic diversity, and between dispersal ability and connectivity. We compiled data on genetic patterns and life history traits in nine species across five phyla. Sampling sites spanned 600 km in the northwest Mediterranean Sea and focused on a 50‐km area near Marseilles, France. Comparative population genetic approaches yielded three main results. (i) Species without larvae showed higher levels of genetic structure than species with free‐living larvae, but the role of larval type (lecithotrophic or planktotrophic) was negligible. (ii) A narrow area around Marseilles, subject to offshore advection, limited genetic connectivity in most species. (iii) We identified sites with significant positive contributions to overall genetic diversity across all species, corresponding with areas near low human population densities. In contrast, high levels of human activity corresponded with a negative contribution to overall genetic diversity. Genetic diversity within species was positively and significantly linearly related to local species diversity. Our study suggests that local contribution to overall genetic diversity should be taken into account for future conservation strategies.     

Dependence of sustainability on the configuration of marine reserves and larval dispersal distance

Marine reserves hold promise for maintaining biodiversity and sustainable fishery management, but studies supporting them have not addressed a crucial aspect of sustainability: the reduction in viability of populations with planktonic larvae dispersing along a coastal habitat with noncontiguous marine reserves. We show how sustainability depends on the fraction of natural larval settlement (FNLS) remaining after reserves are implemented, which in turn depends on reserve configuration and larval dispersal distance. Sustainability requires FNLS to be greater than an empir-ically determined minimum. Maintaining an adequate value for all species requires either a large, unlikely fraction (> 35%) of coastline in reserves, or reserves that are larger than the mean larval dispersal distance of the target species. FNLS is greater for species dispersing shorter distances, which implies reserves can lead to: (1) changes in community composition and (2) genetic selection for shorter dispersal distance. Dependence of sustainability on dispersal distance is a new source of uncertainty.     

Blood cells and coelomocytes of the inarticulate brachiopod Lingula anatina

Three cell types are described from the coelomic cavity of the pedicle of the brachiopod Lingula anatina . Erythrocytes are abundant in the blood vessels of the mantle and also occur, in reduced numbers, in the pedicle. Phagocytic amoebocytes, characterized by a variable number of electrondense, homogeneous granules are common in the coelomic fluid of the pedicle. The enigmatic spindle bodies described by earlier authors constitute the most common cell type encountered in the pedicle coelom of aquarium-maintained specimens. The origin of spindle bodies from muscle cells is suggested.     

Sharp genetic breaks among populations of Haptosquilla pulchella (Stomatopoda) indicate limits to larval transport: patterns, causes, and consequences

To help stem the precipitous decline of coral reef ecosystems world-wide, conservation efforts are focused on establishing interconnected reserve networks to protect threatened populations. Because many coral reef organisms have a planktonic or pelagic larval dispersal phase, it is critical to understand the patterns of ecological connectivity between reserve populations that result from larval dispersal. We used genetics to infer dispersal patterns among 24 Indo-West Pacific populations of the mantis shrimp, Haptosquilla pulchella. Contrary to predictions of high dispersal facilitated by the strong currents of the Indonesian throughflow, mitochondrial DNA sequences from 393 individuals displayed striking patterns of regional genetic differentiation concordant with ocean basins isolated during periods of lowered sea level. Patterns of genetic structuring indicate that although dispersal within geographical regions with semicontiguous coastlines spanning thousands of kilometres may be common, ecologically meaningful connections can be rare among populations separated by as little as 300 km of open ocean. Strong genetic mosaics in a species with high dispersal potential highlight the utility of genetics for identifying regional patterns of genetic connectivity between marine populations and show that the assumption that ocean currents will provide ecological connectivity among marine populations must be empirically tested in the design of marine reserve networks.     

Theoretical limits to the correlation between pelagic larval duration and population genetic structure

Increasing dispersal duration should result in increasing dispersal distance, facilitating higher gene flow among populations. As such, it has long been predicted that genetic structure (e.g. F(ST) ) among populations of marine species should be strongly correlated with pelagic larval duration (PLD). However, previous studies have repeatedly shown a surprisingly poor correspondence. This result has been frequently interpreted as evidence for larval behaviours or physical oceanographic processes that result in larvae failing to reach their dispersal potential, or error inherent in estimating PLD and F(ST) . This study employed a computer modelling approach to explore the impacts of various uncertainties on the correlation between measures of genetic differentiation such as F(ST) and PLD. Results indicate that variation resulting from PLD estimation error had minor impacts on the correlation between genetic structure and PLD. However, variation in effective population size between species, errors in F(ST) estimation and non-equilibrium F(ST) values all had major impacts, resulting in dramatically weaker correlations between PLD and F(ST) . These results suggest that poor correlations between PLD and F(ST) may result from variation and uncertainty in the terms associated with the calculation of F(ST) values. As such, PLD may be a much stronger determinant of realized larval dispersal than suggested by the weak-to-moderate correlations between PLD and F(ST) reported in empirical studies.     

Population structure is not a simple function of reproductive mode and larval type: insights from tropical corals

1. For a wide range of organisms, heritable variation in life-history characteristics has been shown to be strongly subject to selection, reflecting the impact that variation in characters such as genotypic diversity, duration of larval development and adaptations for dispersal can have on the fitness of offspring and the make-up of populations. Indeed, variation in life-history characteristics, especially reproduction and larval type, have often been used to predict patterns of dispersal and resultant population structures in marine invertebrates. 2. Scleractinian corals are excellent models with which to test this relationship, as they exhibit almost every possible combination of reproductive mode and larval type. Some general patterns are emerging but, contrary to expectations, genetic data suggest that while populations of broadcast spawning species may be genotypically diverse they may be heavily reliant on localized recruitment rather than widespread dispersal of larvae. 3. Here we use microsatellites to test the importance of localized recruitment by comparing the genetic structure of populations of two broadcast spawning corals with contrasting modes of reproduction and larval development; Goniastrea favulus is self-compatible, has sticky, negatively buoyant eggs and larvae and is expected to have restricted dispersal of gametes and larvae. In contrast, Platygyra daedalea is self-incompatibile, spawns positively buoyant egg-sperm bundles and has planktonic development. 4. Surprisingly, spatial-autocorrelation revealed no fine-scale clustering of similar genotypes within sites for G. favulus, but showed a non-random distribution of genotypes in P. daedalea. Both species showed similar levels of genetic subdivision among sites separated by 50-100 m (F(ST) = 0.03), suggesting that larval dispersal may be equivalent in both species. 5. Interestingly, as fragmentation has been considered rare in massive corals, our sample of 284 P. daedalea colonies included 28 replicated genotypes that were each unlikely (P < 0.05) to have been derived independently from sexual reproduction. 6. We conclude that the extreme life history of G. favulus does not produce unusually fine-scale genetic structure and subsequently, that reproductive mode and larval type may not be not good predictors of population structure or dispersal ability.     

Coalescent and biophysical models of stepping‐stone gene flow in neritid snails

Marine species in the Indo‐Pacific have ranges that can span thousands of kilometres, yet studies increasingly suggest that mean larval dispersal distances are less than historically assumed. Gene flow across these ranges must therefore rely to some extent on larval dispersal among intermediate ‘stepping‐stone’ populations in combination with long‐distance dispersal far beyond the mean of the dispersal kernel. We evaluate the strength of stepping‐stone dynamics by employing a spatially explicit biophysical model of larval dispersal in the tropical Pacific to construct hypotheses for dispersal pathways. We evaluate these hypotheses with coalescent models of gene flow among high‐island archipelagos in four neritid gastropod species. Two of the species live in the marine intertidal, while the other two are amphidromous, living in fresh water but retaining pelagic dispersal. Dispersal pathways predicted by the biophysical model were strongly favoured in 16 of 18 tests against alternate hypotheses. In regions where connectivity among high‐island archipelagos was predicted as direct, there was no difference in gene flow between marine and amphidromous species. In regions where connectivity was predicted through stepping‐stone atolls only accessible to marine species, gene flow estimates between high‐island archipelagos were significantly higher in marine species. Moreover, one of the marine species showed a significant pattern of isolation by distance consistent with stepping‐stone dynamics. While our results support stepping‐stone dynamics in Indo‐Pacific species, we also see evidence for nonequilibrium processes such as range expansions or rare long‐distance dispersal events. This study couples population genetic and biophysical models to help to shed light on larval dispersal pathways.     

Nucleotide sequence of 5S ribosomal RNA from Lingula anatina. A study on the molecular evolution of 5S ribosomal RNA from a living fossil

By chromatography on columns of DEAE-Sephadex A-50 and Sephadex G-100, and electrophoresis on polyacrylamide gel, 5S rRNA was purified from a low-molecular-weight RNA fraction extracted from the total tissues of Lingula anatina. Complete digests of the 5S rRNA with RNase T1 [EC 3.1.4.8] and pancreatic RNase [EC 3.1.4.22] were sequenced by conventional column chromatography procedures. The nucleotide sequence of this RNA was determined mainly by a chemical method for sequencing the RNA 3' end-labeled with 32P (1), with the complement of the oligonucleotide catalog obtained by the complete RNase digestions of the RNA. By comparing the sequences of several invertebrate, vertebrate, and Chlorella 5S rRNAs, a phylogenic tree of the rRNAs was constructed and the time of divergence of Lingula was estimated.     

Larval Connectivity and the International Management of Fisheries

Predicting the oceanic dispersal of planktonic larvae that connect scattered marine animal populations is difficult, yet crucial for management of species whose movements transcend international boundaries. Using multi-scale biophysical modeling techniques coupled with empirical estimates of larval behavior and gamete production, we predict and empirically verify spatio-temporal patterns of larval supply and describe the Caribbean-wide pattern of larval connectivity for the Caribbean spiny lobster (Panulirus argus), an iconic coral reef species whose commercial value approaches $1 billion USD annually. Our results provide long sought information needed for international cooperation in the management of marine resources by identifying lobster larval connectivity and dispersal pathways throughout the Caribbean. Moreover, we outline how large-scale fishery management could explicitly recognize metapopulation structure by considering larval transport dynamics and pelagic larval sanctuaries.     

Genetic isolation by distance among populations of the netted dog whelk Nassarius reticulatus (L.) along the European Atlantic coastline

Estimates of the average distances by which marine larvae disperse are generally poorly described, despite the central role that larval dispersal plays in the demographic connectivity of populations across geographic space. Here, we describe the population genetic structure and average dispersal distance of the netted dog whelk Nassarius reticulatus (L.) (Mollusca, Gastropoda, Prosobranchia), a widespread member of European intertidal communities, using DNA sequence variation in a fragment of the mitochondrial gene cytochrome c oxidase subunit I (COI). An analysis of 156 individuals from 6 locations spread across approximately 1700 km of the European Atlantic coastline revealed weak and nonsignificant population structure (overall Phi(ST) = 0.00013). However, pairwise Phi(ST) values revealed a slight but significant increase in genetic isolation with geographic distance (IBD), suggesting that populations are not panmictic across the sampled geographic range. If we assume that the isolation by distance is maintained by a stable, stepping stone model of gene flow, then the slope of the IBD is consistent with an average larval dispersal distance of approximately 70 km per generation. The spatial scale of larval dispersal in N. reticulatus is consistent with the life cycle of the species (planktotrophic veliger lasting 30-60 days before competent to settle). A mismatch analysis of the COI sequences revealed a signature of an ancient demographic expansion that began 61 500-160,000 years ago, well before the most recent Pleistocene glaciation event. The greatest levels of genetic diversity occur within the middle latitudes of the whelk's geographic range, consistent with the notion that historic populations of N. reticulatus might have expanded northward and southward from the centrally located Bay of Biscay.     

A modelling study of the respective role of hydrodynamic processes and larval mortality on larval dispersal and recruitment of benthic invertebrates: example of Pectinaria koreni (Annelida: Polychaeta) in the Bay of Seine (English Channel)

The dispersal of Pectinaria koreni larvae released from theeastern Bay of Seine (English Channel) was studied using a two-dimensionalhydrodynamic model which integrates tides, wind-driven currentsand eddy diffusion, in order to examine the influence of environmentalforcing and mortality on larval population dynamics. A broadagreement between predicted larval dispersal for two spawningevents observed in 1987 and field data suggests that numericalmodelling may be useful to analyse processes involved in thetransport and the dynamics of larval populations. Larval mortalitymay be as important as hydrodynamic processes on larval lossesfor the adult population. Tides and eddy diffusion had someeffects on larval dispersal, but wind forcing and the timingof spawning in relation to the meteorological environment arepredicted to be the main source of variability in larval dispersalrates. Although wind-induced larval transport may produce interannualvariations in larval retention and recruitment, predicted retentionrates were always sufficient to ensure the maintenance of theadult population, regardless of hydrodynamic conditions. Thelong-range transport of larvae from the eastern Bay of Seineto distant populations was conditioned by constant strong winds,lasting 15 consecutive days, and should be considered an extremelyrare event.