text:
Sophie Arnaud-Haond
EMAIL : sophie.arnaud ifremer.fr
ADRESSE : Bd Jean Monnet - 34203 - SETE Cedex - FR 2 rue des Chantiers - 34200 - SETE - FR
SITE PERSO : http://annuaire.ifremer.fr/cv/17007/
Institut : /BOUCLE_groupemots> IFREMER
LIEU GEOGRAPHIQUE : /BOUCLE_groupemots> SETE
GRADE : /BOUCLE_groupemots> Chercheur
Thèmes de recherche : /BOUCLE_groupemots> Écologie évolutive et adaptation
Pôles communs : /BOUCLE_groupemots> Expertise
Arnaud-Haond, S., et al. "Entangled fates of holobiont genomes during invasion: nested bacterial and host diversities in Caulerpa taxifolia." Mol. Ecol.. 26.8 (2017): 2379–2391.
Résumé: Successful prevention and mitigation of biological invasions requires retracing the initial steps of introduction, as well as understanding key elements enhancing the adaptability of invasive species. We studied the genetic diversity of the green alga Caulerpa taxifolia and its associated bacterial communities in several areas around the world. The striking congruence of alpha and beta diversity of the algal genome and endophytic communities reveals a tight association, supporting the holobiont concept as best describing the unit of spreading and invasion. Both genomic compartments support the hypotheses of a unique accidental introduction in the Mediterranean and of multiple invasion events in southern Australia. In addition to helping with tracing the origin of invasion, bacterial communities exhibit metabolic functions that can potentially enhance adaptability and competitiveness of the consortium they form with their host. We thus hypothesize that low genetic diversities of both host and symbiont communities may contribute to the recent regression in the Mediterranean, in contrast with the persistence of highly diverse assemblages in southern Australia. This study supports the importance of scaling up from the host to the holobiont for a comprehensive understanding of invasions.
Mots-Clés: Algae; australia; Caulerpa; Chlorophyta; clonal diversity; dna; endophytic communities; genetic diversity; holobiont; invasion paradox; marine invasion; Mediterranean Sea; microsatellite markers; parasites; plant invasions; polymorphism
Arnaud-Haond, S., et al. "Two "pillars" of cold-water coral reefs along Atlantic European margins: Prevalent association of Madrepora oculata with Lophelia pertusa, from reef to colony scale." Deep-Sea Res. Part II-Top. Stud. Oceanogr.. 145 (2017): 110–119.
Résumé: The scleractinian coral Lophelia pertusa has been the focus of deep-sea research since the recognition of the vast extent of coral reefs in North Atlantic waters two decades ago, long after their existence was mentioned by fishermen. These reefs where shown to provide habitat, concentrate biomass and act as feeding or nursery grounds for many species, including those targeted by commercial fisheries. Thus, the attention given to this cold-water coral (CWC) species from researchers and the wider public has increased. Consequently, new research programs triggered research to determine the full extent of the corals geographic distribution and ecological dynamics of "Lophelia reefs". The present study is based on a systematic standardised sampling design to analyze the distribution and coverage of CWC reefs along European margins from the Bay of Biscay to Iceland. Based on Remotely Operated Vehicle (ROV) image analysis, we report an almost systematic occurrence of Madrepora oculata in association with L. pertusa with similar abundances of both species within explored reefs, despite a tendency of increased abundance of L. pertusa compared to M. oculata toward higher latitudes. This systematic association occasionally reached the colony scale, with "twin" colonies of both species often observed growing next to each other when isolated structures were occurring offireefs. Finally, several "false chimaera" were observed within reefs, confirming that colonial structures can be "coral bushes" formed by an accumulation of multiple colonies even at the inter-specific scale, with no need for self-recognition mechanisms. Thus, we underline the importance of the hitherto underexplored M. oculata in the Eastern Atlantic, reestablishing a more balanced view that both species and their yet unknown interactions are required to better elucidate the ecology, dynamics and fate of European CWC reefs in a changing environment.
Mots-Clés: community; bay; Bay of Biscay; Lophelia pertusa; Madrepora oculata; biscay; mediterranean sea; Cold water corals (CWC); deep-sea corals; False-chimaera colonies; fleuve manche; hydrodynamics; Iceland; Ireland; megafauna; mid-norway; ne atlantic
Becheler, R., et al. "Low incidence of clonality in cold water corals revealed through the novel use of a standardized protocol adapted to deep sea sampling." Deep-Sea Res. Part II-Top. Stud. Oceanogr.. 145 (2017): 120–130.
Résumé: Sampling in the deep sea is a technical challenge, which has hindered the acquisition of robust datasets that are necessary to determine the fine-grained biological patterns and processes that may shape genetic diversity. Estimates of the extent of clonality in deep-sea species, despite the importance of clonality in shaping the local dynamics and evolutionary trajectories, have been largely obscured by such limitations. Cold-water coral reefs along European margins are formed mainly by two reef-building species, Lophelia pertusa and Madrepora oculata. Here we present a fine-grained analysis of the genotypic and genetic composition of reefs occurring in the Bay of Biscay, based on an innovative deep-sea sampling protocol. This strategy was designed to be standardized, random, and allowed the georeferencing of all sampled colonies. Clonal lineages discriminated through their Multi-Locus Genotypes (MLG) at 6-7 microsatellite markers could thus be mapped to assess the level of clonality and the spatial spread of clonal lineages. High values of clonal richness were observed for both species across all sites suggesting a limited occurrence of clonality, which likely originated through fragmentation. Additionally, spatial autocorrelation analysis underlined the possible occurrence of fine-grained genetic structure in several populations of both L. pertusa and M. oculata. The two cold-water coral species examined had contrasting patterns of connectivity among canyons, with among-canyon genetic structuring detected in M. oculata, whereas L. pertusa was panmictic at the canyon scale. This study exemplifies that a standardized, random and georeferenced sampling strategy, while challenging, can be applied in the deep sea, and associated benefits outlined here include improved estimates of fine grained patterns of clonality and dispersal that are comparable across sites and among species.
Mots-Clés: diversity; dispersal; disturbance; Lophelia pertusa; Madrepora oculata; organisms; population-structure; asexual reproduction; Clonality; Cold-water coral; Fine-grained spatial genetic structure; gorgonian coral; lophelia-pertusa; pertusa linnaeus 1758; spatial genetic-structure; Standardized sampling
Cowart, D. A., et al. "Investigation of bacterial communities within the digestive organs of the hydrothermal vent shrimp Rimicaris exoculata provide insights into holobiont geographic clustering." PLoS One. 12.3 (2017): e0172543.
Résumé: Prokaryotic communities forming symbiotic relationships with the vent shrimp, Rimicaris exoculata, are well studied components of hydrothermal ecosystems at the Mid-Atlantic Ridge (MAR). Despite the tight link between host and symbiont, the observed lack of spatial genetic structure seen in R. exoculata contrasts with the geographic differentiation detected in specific bacterial ectosymbionts. The geographic clustering of bacterial lineages within a seemingly panmictic host suggests either the presence of finer scale restriction to gene flow not yet detected in the host, horizontal transmission (environmental selection) of its endosymbionts as a consequence of unique vent geochemistry, or vertically transmitted endosymbionts that exhibit genetic differentiation. To identify which hypothesis best fits, we tested whether bacterial assemblages exhibit differentiation across sites or host populations by performing a 16S rRNA metabarcoding survey on R. exoculata digestive prokaryote samples (n = 31) taken from three geochemically distinct vents across MAR: Rainbow, Trans-Atlantic Geotraverse (TAG) and Logatchev. Analysis of communities across two organs (digestive tract, stomach), three molt colors (white, red, black) and three life stages (eggs, juveniles, adults) also provided insights into symbiont transmission mode. Examining both whole communities and operational taxonomic units (OTUs) confirmed the presence of three main epibionts: Epsilonproteobacteria, Mollicutes and Deferribacteres. With these findings, we identified a clear pattern of geographic segregation by vent in OTUs assigned to Epsilonproteobacteria. Additionally, we detected evidence for differentiation among all communities associated to vents and life stages. Overall, results suggest a combination of environmental selection and vertical inheritance of some of the symbiotic lineages.
Mots-Clés: alignment; chamber; deposits; diversity; Ecology; microbial community; mid-atlantic ridge; population; sequence data; sp nov.
Grehan, A. J., et al. "Towards ecosystem based management and monitoring of the deep Mediterranean, North-East Atlantic and Beyond." Deep-Sea Res. Part II-Top. Stud. Oceanogr.. 145 (2017): 1–7.
Mots-Clés: fish; community; ecosystem based management; monitoring; climate change; conservation; fisheries; bering-sea; baltimore; canyons; cold-water coral; deep-water corals; demersal fish; habitat suitability; habitat suitability modelling; litter; norfolk; rockall bank; vulnerable marine ecosystems
Bailleul, D., S. Stoeckel, and S. Arnaud-Haond. "RClone: a package to identify MultiLocus Clonal Lineages and handle clonal data sets in r." Methods Ecol. Evol.. 7.8 (2016): 966–970.
Résumé: Partially, clonal species are common in the Tree of Life. And yet, population genetic models still mostly focus on the extremes: strictly sexual versus purely asexual reproduction. Here, we present an R package built upon genclone software including new functions and several improvements. The RClone package includes functions to handle clonal data sets, allowing (i) checking for data set reliability to discriminate multilocus genotypes (MLGs), (ii) ascertainment of MLG and semi-automatic determination of clonal lineages (MLL), (iii) genotypic richness and evenness indices calculation based on MLGs or MLLs and (iv) describing several spatial components of clonality. RClone allows the one-shot analysis of multipopulation data sets without size limitation, suitable for data sets now increasingly produced through next-generation sequencing. A major improvement compared to existing software is the ability to determine the threshold to cluster similar MLGs into MLLs, based on implemented simulations of sexual events. Several functions allow data importation, conversion and exportation with adegenet, Genetix or Arlequin. RClone is provided with two vignettes to handle analysis on one (RClonequickmanual) or several populations (RCloneqmsevpops).
Mots-Clés: clonal diversity; clonality; clonal population; diversity; genotype; markers; multilocus genotypes; multilocus lineages; organisms; population-genetics; program; software; spatial autocorrelation
BOAVIDA, J., et al. "A Well-Kept Treasure at Depth: Precious Red Coral Rediscovered in Atlantic Deep Coral Gardens (SW Portugal) after 300 Years." Plos One. 11.1 (2016).
Résumé: Background The highly valuable red coral Corallium rubrum is listed in several Mediterranean Conventions for species protection and management since the 1980s. Yet, the lack of data about its Atlantic distribution has hindered its protection there. This culminated in the recent discovery of poaching activities harvesting tens of kg of coral per day from deep rocky reefs off SW Portugal. Red coral was irregularly exploited in Portugal between the 1200s and 1700s, until the fishery collapsed. Its occurrence has not been reported for the last 300 years. Results Here we provide the first description of an Atlantic red coral assemblage, recently rediscovered dwelling at 60-100 m depth in southern Portugal. We report a very slow growth rate (0.23 mm year-1), comparable to Mediterranean specimens. In comparison with most of the Mediterranean reports, the population reaches much larger sizes, estimated to be over one century old, and has a more complex coral branch architecture that promotes a rich assemblage of associated species, with boreal and Mediterranean affinities. Atlantic red coral is genetically distinct, yet mitochondrial analyses suggest that red corals from the Atlantic may have introgressed the Mediterranean ones after migration via the Algeria current. Our underwater surveys, using advanced mixed-gas diving, retrieved lost fishing gear in all coral sites. Besides illegal harvesting, the use and loss of fishing gears, particularly nets, by local fisheries are likely sources of direct impacts on these benthic assemblages. Conclusions We extended the knowledge on the distribution of C. rubrum in the Atlantic, discovered its genetic distinctiveness, and reveal a rich deep-dwelling fauna associated to these coral assemblages. These findings support a barrier role of the Atlantic-Mediterranean transition zone, but reveal also hints of connectivity along its southern margin. The results highlight the genetic and demographic uniqueness of red coral populations from SW Iberia. However, we also report threats to these vulnerable populations by direct and indirect fishing activities and argue that its protection from any mechanically destructive activities is urgent as a precautionary approach. This study advances our understanding of phylogeographic barriers and range edge genetic diversity, and serves as a baseline against which to monitor future human and environmental disturbances to Atlantic C. rubrum.
Dubois, M., et al. "Linking basin-scale connectivity, oceanography and population dynamics for the conservation and management of marine ecosystems." Glob. Ecol. Biogeogr.. 25.5 (2016): 503–515.
Résumé: AimAssessing the spatial structure and dynamics of marine populations is still a major challenge in ecology. The need to manage marine resources from ecosystem and large-scale perspectives is recognized, but our partial understanding of oceanic connectivity limits the implementation of globally pertinent conservation planning. Based on a biophysical model for the entire Mediterranean Sea, this study takes an ecosystem approach to connectivity and provides a systematic characterization of broad-scale larval dispersal patterns. It builds on our knowledge of population dynamics and discusses the ecological and management implications. LocationThe semi-enclosed Mediterranean Sea and its marine ecosystems are used as a case study to investigate broad-scale connectivity patterns and to relate them to oceanography and population dynamics. MethodsA flow network is constructed by evenly subdividing the basin into sub-regions which are interconnected through the transport of larvae by ocean currents. It allows for the computation of various connectivity metrics required to evaluate larval retention and exchange. ResultsOur basin-scale model predicts that retention processes are weak in the open ocean while they are significant in the coastal ocean and are favoured along certain coastlines due to specific oceanographic features. Moreover, we show that wind-driven divergent (convergent, respectively) oceanic regions are systematically characterized by larval sources (sinks, respectively). Finally, although these connectivity metrics have often been studied separately in the literature, we demonstrate they are interrelated under particular conditions. Their integrated analysis facilitates the appraisal of population dynamics, informing both genetic and demographic connectivities. Main conclusionsThis modelling framework helps ecologists and geneticists to formulate improved hypotheses of population structures and gene flow patterns and to design their sampling strategy accordingly. It is also useful in the implementation and assessment of future protection strategies, such as coastal and offshore marine reserves, by accounting for large-scale dispersal patterns, a missing component of current ecosystem management.
Mots-Clés: coral-reef fish; dispersal; genetic-structure; Larval dispersal; local retention; local retention; marine connectivity; marine ecosystems; marine protected areas; mediterranean littoral fishes; Mediterranean Sea; metapopulation; pelagic larval duration; population dynamics; Population Genetics; protected-area design; sea; self-recruitment; sink dynamics; source
Meistertzheim, A. - L., et al. "Patterns of bacteria-host associations suggest different ecological strategies between two reef building cold-water coral species." Deep Sea Research Part I: Oceanographic Research Papers. 114 (2016): 12–22.
Résumé: Cold-water corals (CWC) are main ecosystem engineers of the deep sea, and their reefs constitute hot-spots of biodiversity. However, their ecology remains poorly understood, particularly, the nature of the holobiont formed by corals with their associated bacterial communities. Here, we analyzed Madrepora oculata and Lophelia pertusa samples, collected from one location in a Mediterranean canyon in two different seasons (autumn and spring), in order to test for species specificity and temporal stability of the host-bacteria associations. The 16S rRNA sequencing revealed host-specific patterns of bacterial communities associated with L. pertusa and M. oculata, both in terms of community composition and diversity. All analyzed M. oculata polyps exhibited temporally and spatially similar bacterial communities dominated by haplotypes homologous to the known cnidarians-associated genus Endozoicomonas. In contrast, the bacterial communities associated with L. pertusa varied among polyps from the same colony, as well as among distinct colonies and between seasons. While the resilient consortium formed by M. oculata and its bacterial community fit the definition of holobiont, the versatility of the L. pertusa microbiome suggests that this association is more influenced by the environmental conditions or nutritional status. Our results thus highlight distinct host/microbes association strategies for these two closely related Scleractinians sharing the same habitat, suggesting distinct sensitivity to environmental change.
Mots-Clés: Bacterial communities; Bacterial ecology; Deep-Sea corals; Lophelia pertusa; Madrepora oculata; Mediterranean Sea; Microbe-microbe and microbe-host interactions
RANJBAR, M. S., et al. "Rising the Persian Gulf Black-Lip Pearl Oyster to the Species Level: Fragmented Habitat and Chaotic Genetic Patchiness in Pinctada persica." Evolutionary Biology. 43.1 (2016): 131–143.
Résumé: Marine organisms with long pelagic larval stages are expected to exhibit low genetic differentiation due to their potential to disperse over large distances. Growing body of evidence, however, suggests that marine populations can differentiate over small spatial scales. Here we focused on black-lip pearl oysters from the Persian Gulf that are thought to belong to the Pinctada margaritifera complex given their morphological affinities. This species complex includes seven lineages that show a wide distribution ranging from the Persian Gulf (Pinctada margaritifera persica) and Indian Ocean (P. m. zanzibarensis) to the French Polynesia (P. m. cumingii) and Hawai'i (P. m. galtsoffi). Despite the long pelagic larval phase of P. m. persica, this lineage is absent from continental locations and can only be found on a few islands of the Persian Gulf. Mitochondrial COI-based analyses indicated that P. m. persica belongs to a clearly divergent ESU and groups with specimens from Mauritius (P. m. zanzibarensis). Microsatellite data, used here to assess the spatial scale of realized dispersal of Persian Gulf black-lip pearl oysters, revealed significant genetic structure among islands distant of only a few dozen kilometres. The scantiness of suitable habitats most likely restricted the distribution of this lineage originating the observed chaotic genetic patchiness. The hatchery-based enhancement performed in one of the sampled islands may also have affected population genetic structure. The long-term accumulation of genetic differences likely resulted from the allopatric divergence between P. m. persica and the neighbouring Indian Ocean black-lip pearl oysters.
Mots-Clés: Black-lip pearl oysters; Fragmented habitat; Persian Gulf; Pinctada persica; Species delimitation
Reichel, K., et al. "Rare sex or out of reach equilibrium? The dynamics of F-IS in partially clonal organisms." BMC Genet.. 17 (2016): 76.
Résumé: Background: Partially clonal organisms are very common in nature, yet the influence of partial asexuality on the temporal dynamics of genetic diversity remains poorly understood. Mathematical models accounting for clonality predict deviations only for extremely rare sex and only towards mean inbreeding coefficient (F-IS) over bar < 0. Yet in partially clonal species, both F-IS < 0 and F-IS > 0 are frequently observed also in populations where there is evidence for a significant amount of sexual reproduction. Here, we studied the joint effects of partial clonality, mutation and genetic drift with a state-and-time discrete Markov chain model to describe the dynamics of F-IS over time under increasing rates of clonality. Results: Results of the mathematical model and simulations show that partial clonality slows down the asymptotic convergence to F-IS = 0. Thus, although clonality alone does not lead to departures from Hardy-Weinberg expectations once reached the final equilibrium state, both negative and positive F-IS values can arise transiently even at intermediate rates of clonality. More importantly, such "transient" departures from Hardy Weinberg proportions may last long as clonality tunes up the temporal variation of F-IS and reduces its rate of change over time, leading to a hyperbolic increase of the maximal time needed to reach the final mean (F-IS,F-infinity) over bar value expected at equilibrium. Conclusion: Our results argue for a dynamical interpretation of F-IS in clonal populations. Negative values cannot be interpreted as unequivocal evidence for extremely scarce sex but also as intermediate rates of clonality in finite populations. Complementary observations (e.g. frequency distribution of multiloci genotypes, population history) or time series data may help to discriminate between different possible conclusions on the extent of clonality when mean (F-IS) over bar values deviating from zero and/or a large variation of F-IS over loci are observed.
Mots-Clés: consequences; diversity; evolution; genetic diversity; genetic-structure; Heterozygote excess; hybridization; Inbreeding coefficient; loci; Mating system; microsatellite primers; mutation; Parthenogenesis; Partial asexuality; population-genetics; reproduction
Aires, T., et al. "Hologenome theory supported by cooccurrence networks of species-specific bacterial communities in siphonous algae (Caulerpa)." FEMS Microbiology Ecology. 91.7 (2015): fiv067.
Résumé: The siphonous algae of the Caulerpa genus harbor internal microbial communities hypothesized to play important roles in development, defense and metabolic activities of the host. Here, we characterize the endophytic bacterial community of four Caulerpa taxa in the Mediterranean Sea, through 16S rRNA amplicon sequencing. Results reveal a striking alpha diversity of the bacterial communities, similar to levels found in sponges and coral holobionts. These comprise (1) a very small core community shared across all hosts (< 1% of the total community), (2) a variable portion (ca. 25%) shared by some Caulerpa taxa but not by all, which might represent environmentally acquired bacteria and (3) a large (>70%) species-specific fraction of the community, forming very specific clusters revealed by modularity in networks of cooccurrence, even in areas where distinct Caulerpa taxa occurred in sympatry. Indirect inferences based on sequence homology suggest that these communities may play an important role in the metabolism of their host, in particular on their ability to grow on anoxic sediment. These findings support the hologenome theory and the need for a holistic framework in ecological and evolutionary studies of these holobionts that frequently become invasive.
COWART, D. A., et al. "Metabarcoding is powerful yet still blind: a comparative analysis of morphological and molecular surveys of seagrass communities." Plos One. 10.2 (2015): 1–26.
Résumé: In the context of the sixth wave of extinction, reliable surveys of biodiversity are increasingly needed to infer the cause and consequences of species and community declines, identify early warning indicators of tipping points, and provide reliable impact assessments before engaging in activities with potential environmental hazards. DNA metabarcoding has emerged as having potential to provide speedy assessment of community structure from environmental samples. Here we tested the reliability of metabarcoding by comparing morphological and molecular inventories of invertebrate communities associated with sea-grasses through estimates of alpha and beta diversity, as well as the identification of the most abundant taxa. Sediment samples were collected from six Zostera marina seagrass meadows across Brittany, France. Metabarcoding surveys were performed using both mitochondrial (Cytochrome Oxidase I) and nuclear (small subunit 18S ribosomal RNA) markers, and compared to morphological inventories compiled by a long-term benthic monitoring network. A sampling strategy was defined to enhance performance and accuracy of results by preventing the dominance of larger animals, boosting statistical support through replicates, and using two genes to compensate for taxonomic biases. Molecular barcodes proved powerful by revealing a remarkable level of diversity that vastly exceeded the morphological survey, while both surveys identified congruent differentiation of the meadows. However, despite the addition of individual barcodes of common species into taxonomic reference databases, the retrieval of only 36% of these species suggest that the remaining were either not present in the molecular samples or not detected by the molecular screening. This finding exemplifies the necessity of comprehensive and well-curated taxonomic reference libraries and multi-gene surveys. Overall, results offer methodological guidelines and support for metabarcoding as a powerful and repeatable method of characterizing communities, while also presenting suggestions for improvement, including implementation of pilotstudies prior to performing full "blind" metabarcoding assessments to optimize sampling and amplification protocols.
Durand, L., et al. "Biogeographical distribution of Rimicaris exoculata resident gut epibiont communities along the Mid-Atlantic Ridge hydrothermal vent sites." FEMS Microbiology Ecology. 91.10 (2015): fiv101.
Résumé: Rimicaris exoculata is a deep-sea hydrothermal vent shrimp whose enlarged gill chamber houses a complex trophic epibiotic community. Its gut harbours an autochthonous and distinct microbial community. This species dominates hydrothermal ecosystem megafauna along the Mid-Atlantic Ridge, regardless of contrasting geochemical conditions prevailing in them. Here, the resident gut epibiont community at four contrasted hydrothermal vent sites (Rainbow, TAG, Logatchev and Ashadze) was analysed and compiled with previous data to evaluate the possible influence of site location, using 16S rRNA surveys and microscopic observations (transmission electron microscopy, scanning electron microscopy and fluorescence in situ hybridization analyses). Filamentous epibionts inserted between the epithelial cell microvilli were observed on all examined samples. Results confirmed resident gut community affiliation to Deferribacteres, Mollicutes, Epsilonproteobacteria and to a lesser extent Gammaproteobacteria lineages. Still a single Deferribacteres phylotype was retrieved at all sites. Four Mollicutes-related operational taxonomic units were distinguished, one being only identified on Rainbow specimens. The topology of ribotype median-joining networks illustrated a community diversification possibly following demographic expansions, suggesting a more ancient evolutionary history and/or a larger effective population size at Rainbow. Finally, the gill chamber community distribution was also analysed through ribotype networks based on sequences from R. exoculata collected at the Rainbow, Snake Pit, TAG, Logatchev and Ashadze sites. Results allow the refining of hypotheses on the epibiont role and transmission pathways.
GAGNAIRE, P. - A., et al. "Using neutral, selected, and hitchhiker loci to assess connectivity of marine populations in the genomic era." Evolutionary Applications. 8.8 (2015): 769–786.
Résumé: Estimating the rate of exchange of individuals among populations is a central concern to evolutionary ecology and its applications to conservation and management. For instance, the efficiency of protected areas in sustaining locally endangered populations and ecosystems depends on reserve network connectivity. The population genetics theory offers a powerful framework for estimating dispersal distances and migration rates from molecular data. In the marine realm, however, decades of molecular studies have met limited success in inferring genetic connectivity, due to the frequent lack of spatial genetic structure in species exhibiting high fecundity and dispersal capabilities. This is especially true within biogeographic regions bounded by well-known hotspots of genetic differentiation. Here, we provide an overview of the current methods for estimating genetic connectivity using molecular markers and propose several directions for improving existing approaches using large population genomic datasets. We highlight several issues that limit the effectiveness of methods based on neutral markers when there is virtually no genetic differentiation among samples. We then focus on alternative methods based on markers influenced by selection. Although some of these methodologies are still underexplored, our aim was to stimulate new research to test how broadly they are applicable to nonmodel marine species. We argue that the increased ability to apply the concepts of cline analyses will improve dispersal inferences across physical and ecological barriers that reduce connectivity locally. We finally present how neutral markers hitchhiking with selected loci can also provide information about connectivity patterns within apparently well-mixed biogeographic regions. We contend that one of the most promising applications of population genomics is the use of outlier loci to delineate relevant conservation units and related eco-geographic features across which connectivity can be measured.
KIVELA, M., S. ARNAUD-HAOND, and J. SARAMAKI. "EDENetworks: A user-friendly software to build and analyse networks in biogeography, ecology and population genetics." Molecular Ecology Resources. 15.1 (2015): 117–122.
Résumé: The recent application of graph-based network theory analysis to biogeography, community ecology and population genetics has created a need for user-friendly software, which would allow a wider accessibility to and adaptation of these methods. EDENetworks aims to fill this void by providing an easy-to-use interface for the whole analysis pipeline of ecological and evolutionary networks starting from matrices of species distributions, genotypes, bacterial OTUs or populations characterized genetically. The user can choose between several different ecological distance metrics, such as Bray-Curtis or Sorensen distance, or population genetic metrics such as FST or Goldstein distances, to turn the raw data into a distance/dissimilarity matrix. This matrix is then transformed into a network by manual or automatic thresholding based on percolation theory or by building the minimum spanning tree. The networks can be visualized along with auxiliary data and analysed with various metrics such as degree, clustering coefficient, assortativity and betweenness centrality. The statistical significance of the results can be estimated either by resampling the original biological data or by null models based on permutations of the data.
Mots-Clés: Biogeography; biological communities; graph theory; microbial ecology; network analysis; population genetics
PANTE, E., et al. "Species are hypotheses: avoid connectivity assessments based on pillars of sand." Molecular Ecology. 24.3 (2015): 525–544.
Résumé: Connectivity among populations determines the dynamics and evolution of populations, and its assessment is essential in ecology in general and in conservation biology in particular. The robust basis of any ecological study is the accurate delimitation of evolutionary units, such as populations, metapopulations and species. Yet a disconnect still persists between the work of taxonomists describing species as working hypotheses and the use of species delimitation by molecular ecologists interested in describing patterns of gene flow. This problem is particularly acute in the marine environment where the inventory of biodiversity is relatively delayed, while for the past two decades, molecular studies have shown a high prevalence of cryptic species. In this study, we illustrate, based on marine case studies, how the failure to recognize boundaries of evolutionary-relevant unit leads to heavily biased estimates of connectivity. We review the conceptual framework within which species delimitation can be formalized as falsifiable hypotheses and show how connectivity studies can feed integrative taxonomic work and vice versa. Finally, we suggest strategies for spatial, temporal and phylogenetic sampling to reduce the probability of inadequately delimiting evolutionary units when engaging in connectivity studies.
Mots-Clés: connectivity; marine organisms; molecular systematics; taxonomy
ADJEROUD, M., et al. "Genetic diversity, clonality and connectivity in the scleractinian coral Pocillopora damicornis: a multi-scale analysis in an insular, fragmented reef system." Marine Biology. 161.3 (2014): 531–541.
Résumé: Clonality and genetic structure of the coral Pocillopora damicornis sensu lato were assessed using five microsatellites in 12 populations from four islands of the Society Archipelago (French Polynesia) sampled in June 2008. The 427 analysed specimens fell into 132 multilocus genotypes (MLGs), suggesting that asexual reproduction plays an important role in the maintenance of these populations. A haploweb analysis of ITS2 sequences of each MLG was consistent with all of them being conspecific. Genetic differentiation was detected both between and within islands, but when a single sample per MLG was included in the analyses, the populations turned out to be nearly panmictic. These observations provide further evidence of the marked variability in reproductive strategies and genetic structure of P. damicornis throughout its geographic range; comparison with results previously obtained for the congeneric species Pocillopora meandrina underlines the importance of life history traits in shaping the genetic structure of coral populations
Arnaud-Haond, S., et al. "Discriminating Micropathogen Lineages and Their Reticulate Evolution through Graph Theory-Based Network Analysis: The Case of Trypanosoma cruzi, the Agent of Chagas Disease." PLoS ONE. 9.8 (2014).
Résumé: Micropathogens (viruses, bacteria, fungi, parasitic protozoa) share a common trait, which is partial clonality, with wide variance in the respective influence of clonality and sexual recombination on the dynamics and evolution of taxa. The discrimination of distinct lineages and the reconstruction of their phylogenetic history are key information to infer their biomedical properties. However, the phylogenetic picture is often clouded by occasional events of recombination across divergent lineages, limiting the relevance of classical phylogenetic analysis and dichotomic trees. We have applied a network analysis based on graph theory to illustrate the relationships among genotypes of Trypanosoma cruzi, the parasitic protozoan responsible for Chagas disease, to identify major lineages and to unravel their past history of divergence and possible recombination events. At the scale of T. cruzi subspecific diversity, graph theory-based networks applied to 22 isoenzyme loci (262 distinct Multi-Locus-Enzyme-Electrophoresis -MLEE) and 19 microsatellite loci (66 Multi-Locus-Genotypes -MLG) fully confirms the high clustering of genotypes into major lineages or "near-clades". The release of the dichotomic constraint associated with phylogenetic reconstruction usually applied to Multilocus data allows identifying putative hybrids and their parental lineages. Reticulate topology suggests a slightly different history for some of the main "near-clades", and a possibly more complex origin for the putative hybrids than hitherto proposed. Finally the sub-network of the near-clade T. cruzi I (28 MLG) shows a clustering subdivision into three differentiated lesser near-clades ("Russian doll pattern"), which confirms the hypothesis recently proposed by other investigators. The present study broadens and clarifies the hypotheses previously obtained from classical markers on the same sets of data, which demonstrates the added value of this approach. This underlines the potential of graph theory-based network analysis for describing the nature and relationships of major pathogens, thereby opening stimulating prospects to unravel the organization, dynamics and history of major micropathogen lineages.
ARNAUD-HAOND, S., et al. "Disentangling the Influence of Mutation and Migration in Clonal Seagrasses Using the Genetic Diversity Spectrum for Microsatellites." Journal Of Heredity. 105.4 (2014): 532–541.
Résumé: The recurrent lack of isolation by distance reported at regional scale in seagrass species was recently suggested to stem from stochastic events of large-scale dispersal. We explored the usefulness of phylogenetic information contained in microsatellite loci to test this hypothesis by using the Genetic Diversity Spectrum (GDS) on databases containing, respectively, 7 and 9 microsatellites genotypes for 1541 sampling units of Posidonia oceanica and 1647 of Cymodocea nodosa. The simultaneous increase of microsatellite and geographic distances that emerges reveals a coherent pattern of isolation by distance in contrast to the chaotic pattern previously described using allele frequencies, in particular, for the long-lived P. oceanica. These results suggest that the lack of isolation by distance, rather than the resulting from rare events of large-scale dispersal, reflects at least for some species a stronger influence of mutation over migration at the scale of the distribution range. The global distribution of genetic polymorphism may, therefore, result predominantly from ancient events of step-by-step (re)colonization followed by local recruitment and clonal growth, rather than contemporary gene flow. The analysis of GDS appears useful to unravel the evolutionary forces influencing the dynamics and evolution at distinct temporal and spatial scales by accounting for phylogenetic information borne by microsatellites, under an appropriate mutation model. This finding adds nuance to the generalization of the influence of large-scale dispersal on the dynamics of seagrasses.
Mots-Clés: clonality; genetic divergence; Genetic Diversity Spectrum; microsatellites; Seagrass; stepwise mutation
BECHELER, R., et al. "Scaling of processes shaping the clonal dynamics and genetic mosaic of seagrasses through temporal genetic monitoring." Heredity. 112.2 (2014): 114–121.
Résumé: Theoretically, the dynamics of clonal and genetic diversities of clonal plant populations are strongly influenced by the competition among clones and rate of seedling recruitment, but little empirical assessment has been made of such dynamics through temporal genetic surveys. We aimed to quantify 3 years of evolution in the clonal and genetic composition of Zostera marina meadows, comparing parameters describing clonal architecture and genetic diversity at nine microsatellite markers. Variations in clonal structure revealed a decrease in the evenness of ramet distribution among genets. This illustrates the increasing dominance of some clonal lineages (multilocus lineages, MLLs) in populations. Despite the persistence of these MLLs over time, genetic differentiation was much stronger in time than in space, at the local scale. Contrastingly with the short-term evolution of clonal architecture, the patterns of genetic structure and genetic diversity sensu stricto (that is, heterozygosity and allelic richness) were stable in time. These results suggest the coexistence of (i) a fine grained (at the scale of a 20 x 30 m quadrat) stable core of persistent genets originating from an initial seedling recruitment and developing spatial dominance through clonal elongation; and (ii) a local (at the scale of the meadow) pool of transient genets subjected to annual turnover. This simultaneous occurrence of initial and repeated recruitment strategies highlights the different spatial scales at which distinct evolutionary drivers and mating systems (clonal competition, clonal growth, propagule dispersal and so on) operate to shape the dynamics of populations and the evolution of polymorphism in space and time.
Mots-Clés: clonality; Seagrass; spatio-temporal genetic structure; Zostera marina
BROGGIATO, A., et al. "Fair and equitable sharing of benefits from the utilization of marine genetic resources in areas beyond national jurisdiction: Bridging the gaps between science and policy." Marine Policy. 49 (2014): 176–185.
Résumé: Marine genetic resources are a subject of a growing body of research and development activities, as demonstrated by the abundance of marine patented genes reported in GenBank. Given the lack of a comprehensive legal regime for the management of marine genetic resources in areas beyond national jurisdiction, the General Assembly of the United Nations met in 2006 to discuss whether there are regulatory or governance gaps and how to address them. Besides the crystallization of the different political positions, the process is now advancing towards making a decision about whether to develop an international instrument under the United Nations Convention on the Law of the Sea (UNCLOS) for the conservation and sustainable use of marine biological diversity, within which the regulation of access to genetic resources and the sharing of benefits from their utilization has emerged as an in-dissociable issue. In order to propose concrete options to be considered for the establishment of a legal framework addressing these issues, policy-makers need to better understand the feasibility, the costs and the modalities of scientific activities undertaken, together with the actual level of commercialization of new products. They also need to be aware of the already advanced practices in place within the scientific community, especially regarding sharing of non-monetary benefits. This paper particularly highlights and discusses practical scenarios to advance in the international process, based on the approaches adopted in other regional and international regimes for the management of genetic resources and on the best practices developed within the scientific community.
Mots-Clés: Access and benefit-sharing; Common pool resources; Intellectual property rights; Marine genetic resources; Marine scientific research; Public domain
LAGABRIELLE, E., et al. "Connecting MPAs – eight challenges for science and management." Aquatic Conservation-marine And Freshwater Ecosystems. 24 (2014): 94–110.
Résumé: Connectivity is a crucial process underpinning the persistence, recovery, and productivity of marine ecosystems. The Convention on Biological Diversity, through the Aichi Target 11, has set the ambitious objective of implementing a 'well connected system of protected areas' by 2020. This paper identifies eight challenges toward the integration of connectivity into MPA network management and planning. A summary table lists the main recommendations in terms of method, tool, advice, or action to address each of these challenges. Authors belong to a science–management continuum including researchers, international NGO officers, and national MPA agency members. Three knowledge challenges are addressed: selecting and integrating connectivity measurement metrics; assessing the accuracy and uncertainty of connectivity measurements; and communicating and visualizing connectivity measurements. Three management challenges are described: integrating connectivity into the planning and management of MPA networks; setting quantitative connectivity targets; and implementing connectivity-based management across scales and marine jurisdictions. Finally, two paths toward a better integration of connectivity science with MPA management are proposed: setting management-driven priorities for connectivity research, bridging connectivity science, and MPA network management. There is no single method to integrate connectivity into marine spatial planning. Rather, an array of methods can be assembled according to the MPA network objectives, budget, available skills, data, and timeframe. Overall, setting up 'boundary organizations' should be promoted to organize complex cross-disciplinary, cross-sectoral and cross-jurisdiction interactions that are needed between scientists, managers, stakeholders and decision-makers to make informed decision regarding connectivity-based MPA planning and management.
Mots-Clés: coastal; conservation evaluation; Fishing; Marine Protected Areas; ocean; spatial modelling
Van Dover, C. L., et al. "Ecological Restoration in the Deep Sea: Desiderata." Marine Policy. 44 (2014): 98–106.
Résumé: An era of expanding deep-ocean industrialization is before us, with policy makers establishing governance frameworks for sustainable management of deep-sea resources while scientists learn more about the ecological structure and functioning of the largest biome on the planet. Missing from discussion of the stewardship of the deep ocean is ecological restoration. If existing activities in the deep sea continue or are expanded and new deep-ocean industries are developed, there is need to consider what is required to minimize or repair resulting damages to the deep-sea environment. In addition, thought should be given as to how any past damage can be rectified. This paper develops the discourse on deep-sea restoration and offers guidance on planning and implementing ecological restoration projects for deep-sea ecosystems that are already, or are at threat of becoming, degraded, damaged or destroyed. Two deep-sea restoration case studies or scenarios are described (deep-sea stony corals on the Darwin Mounds off the west coast of Scotland, deep-sea hydrothermal vents in Manus Basin, Papua New Guinea) and are contrasted with on-going saltmarsh restoration in San Francisco Bay. For these case studies, a set of socio-economic, ecological, and technological decision parameters that might favor (or not) their restoration are examined. Costs for hypothetical restoration scenarios in the deep sea are estimated and first indications suggest they may be two to three orders of magnitude greater per hectare than costs for restoration efforts in shallow-water marine systems.
Mots-Clés: Cold-water corals; Deep-sea resource use; Hydrothermal vents; Marine policy; Restoration science
|Aires, T., et al. "Invasion Is a Community Affair: Clandestine Followers in the Bacterial Community Associated to Green Algae, Caulerpa racemosa, Track the Invasion Source." PLoS ONE. 8.7 (2013).
|Arnaud-Haond, S., et al. "Microsatellite markers developed through pyrosequencing allow clonal discrimination in the invasive alga Caulerpa taxifolia." Conservation Genetics Resources. 5.3 (2013): 667–669.
|Gérard, K., et al. "PCR survey of 50 introns in animals: Cross-amplification of homologous EPIC loci in eight non-bilaterian, protostome and deuterostome phyla." Marine Genomics. 12 (2013): 1–8.
|Massa, S. I., et al. "Entangled effects of allelic and clonal (genotypic) richness in the resistance and resilience of experimental populations of the seagrass Zostera noltii to diatom invasion." BMC Ecology. 13.1 (2013).
|Meistertzheim, A. - L., et al. "Genetic structure of wild European populations of the invasive Pacific oyster Crassostrea gigas due to aquaculture practices." Marine Biology. 160.2 (2012): 453–463.


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Now try the following tasks. If you have problems completing them, contact your tutor
Task 1
Using a paper-based approach work your way through at least one example of Shell sort. You may wish to use the data set used here initially, but you should try at least one data set of your own making.
Task 2
Implement a function called shellsort to be used with the program you developed for previous sections. You will find that you will need to pay great care with loop counters for this exercise. For this first implementation you should use slice sizes of 2, 4, 8 and so on until it reaches the insertion sort stage. Count the number of comparisons and moves required to sort the datasets.
Task 3
Implement a function called shellsort2. This should differ from the previous version in that it has a second array embedded in the function that holds the number of slices to be used for a given pass. (In the bench-checked example above the values in the array would start 3, 5 and 10)
Use this function to experiment with different sequences of slices (in general the program uses a sequence in increasing order) . Record the number of comparisons and moves required to sort the datasets for different sequences.
Next: Merge Sort

subset: commoncrawl


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Price: $10.00
Quantity: 1 available
Very good copy of first paperback printing 1963, first thus. 156 pp. One three-act and four one-act plays. Cover rubbed and creased, spine sunned, spine slanted.
Title: Lost Plays of Eugene O'Neill, 1913-1915
Categories: Other,
Edition: First Paperbound Edition
Publisher: New York, Citadel Press: 1963
Binding: Wraps
Book Condition: Very Good
Type: Wraps
Item: 1.00 Item
Seller ID: 000255
Keywords: Drama


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'Tolled' means 'stop the clock' for time to refile state-law claims after federal claims dismissed
In a procedural decision arising out of dismissed federal and state employment discrimination claims, a 5-4 Supreme Court held that 28 U.S.C. §1367(d), which says that the period of limitations for refiling state claims after they have been dismissed in a federal court action "shall be tolled while the claim is pending [in federal court]," stops the clock while the claim is in federal court, and it starts running again when the tolling period ends. Accordingly, an employee's state-law claims that were dismissed from federal court after two and a half years and were refiled in state court 59 days later were timely, because the applicable statute of limitations for her D.C. law claims was suspended while her federal suit was pending. Notably, Chief Justice Roberts joined the liberal wing of the court in the majority opinion authored by Justice Ginsburg; Justice Gorsuch dissented (Artis v. District of Columbia, January 22, 2018, Ginsburg, R.).
What to do with dismissed state-law claims? Federal district courts may exercise supplemental jurisdiction over state claims not otherwise within their adjudicatory authority if those claims are "part of the same case or controversy" as the federal claims the plaintiff asserts. When a district court dismisses all claims independently qualifying for the exercise of federal jurisdiction, it ordinarily also dismisses all related state claims. 28 U.S.C. §1367(d) provides that the "period of limitations for" refiling in state court a state claim so dismissed "shall be tolled while the claim is pending [in federal court] and for a period of 30 days after it is dismissed unless State law provides for a longer tolling period."
When the employee, a health inspector, filed a federal-court suit against the District of Columbia alleging a federal Title VII employment-discrimination claim and three allied claims under D.C. law, nearly two years remained on the applicable statute of limitations for her D.C.-law violations, which included whistleblower retaliation, statutory wrongful termination, and public policy wrongful termination. Two and a half years later, the district court ruled against the employee on her sole federal claim and dismissed the D.C.-law claims; 59 days after that dismissal, the employee refiled her state-law claims in the D.C. state court, but that court dismissed them as time-barred. The appeals court affirmed, holding that the statute gave the employee only a 30-day grace period to refile in state court because that "better accommodates federalism concerns"; it rejected her argument that "tolled" in Section 1367(d) means that the limitations period was suspended during the pendency of the federal suit.
The Supreme Court majority framed the issue to be whether "the word 'tolled,' as used in §1367(d), means the state limitations period is suspended during the pendency of the federal suit; or does 'tolled' mean that, although the state limitations period continues to run, a plaintiff is accorded a grace period of 30 days to refile in state court post dismissal of the federal case?"
To "toll" means to stop the clock. According to the majority's reasoning, there are two ways that statutes protecting claims that were filed in another forum from being time-barred work. One is to be "tolled" or suspended, while the claim is pending elsewhere; "the time clock starts running again when the tolling period ends, picking up where it left off." Or a legislature may elect to provide a grace period, allowing the statute of limitations to run while the claim is pending in another forum and giving the plaintiff a fixed period in which to refile.
The statute uses "toll." Section 1367(d) uses the language of a tolling provision: "It suspends the statute of limitations both while the claim is pending in federal court and for 30 days post dismissal," said the majority, agreeing with the employee's interpretation. The District, in contrast, had read "tolled" to mean to remove, temporarily, the bar that would ordinarily accompany the expiration of the limitations period, which the Court said was "a strained interpretation of the phrase 'period of limitations' and 'could yield an absurdity, permitting a plaintiff to refile in state court even if the limitations period on her claim had expired before she filed in federal court.'"
It was significant to the High Court that the District had not identified any federal statute in which a grace-period meaning has been ascribed to the word "tolled" or any word similarly rooted. Nor had the dissent "identified even one federal statute that fits its bill, i.e., a federal statute that says 'tolled' but means something other than 'suspended,' or 'paused,' or 'stopped.'"
Plus a grace period. After disposing of the argument that Congress had adopted an American Law Institute 1969 recommendation to allow refiling in state court only for 30 days after a dismissal, the majority turned to Section 1367(d)'s 30-day federal floor on the time allowed for refiling in state court. Why would that grace-period provision be included, suggested the District, if the tolling provision meant suspend or stop the clock? This additional 30-day grace period accounted for cases in which a plaintiff filed a federal action close to the date the relevant state statute of limitations expired, by giving a plaintiff "breathing space" to refile in state court, reasoned the Court, finding it not unusual in stop-the-clock statutes. The Court also noted Section 1367(d)'s proviso "unless State law provides for a longer tolling period," which also would aid a plaintiff who filed in federal court just before the state limitations period was set to expire.
Exceed Congressional authority? Although the District also asked whether the statute exceeded Congress' authority under the Necessary and Proper Clause "because its connection to Congress' enumerated powers is too attenuated or because it is too great an incursion on the States' domain," the majority found that a "stop-the-clock" interpretation of Section 1367(d) did not present a serious constitutional problem. Specifically, in Jinks v. Richland County (2003), the Court earlier rejected an argument that Section 1367(d) impermissibly exceeded Congress' authority under the Necessary and Proper Clause. The District contended that a stop-the-clock prescription serves "no federal purpose" that could not be served by a grace-period prescription. Both a stop-the-clock prescription and a grace-period prescription are "standard, off-the-shelf means of accounting for the fact that a claim was timely" in another forum, reasoned the Court.
"Requiring Congress to choose one over the other would impose a tighter constraint on Congress' discretion than this Court has countenanced," and concerns that a stop-the-clock approach is a greater imposition on the states than a grace-period approach "may also be more theoretical than real," the majority concluded, reversing and remanding. A stop-the-clock rule supports primary reason behind limitations statutes: "preventing surprises" to defendants and "barring a plaintiff who has slept on his rights," continued the majority, because when the statute applies, defendants will have notice of the plaintiff's claims within the state-prescribed limitations period as a plaintiff will have timely asserted those rights by attempting to pursue them in one litigation in federal court.
Dissent. Justice Gorsuch, although acknowledging early on that the majority opinion's interpretation of "toll" was both possible and historically common, predicted the majority's approach "will require state courts to entertain state law claims that state law deems untimely not only by weeks or months but by many years." He went on to explain that "the Court demands all this without offering any rational account why it is necessary or proper to the exercise of one of the federal government's limited and enumerated powers. It may only be a small statute we are interpreting, but the result the Court reaches today represents no small intrusion on traditional state functions and no small departure from our foundational principles of federalism."


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a gravelly beach
- a gravelly beach. 碎石滩
- "This stuff used to be illegal," he says in a gravelly voice. "这些东西原本是违法的",他用沙哑的声音说。
- The bows of the canoe grounded upon the gravelly beach with a gentle motion, and a sound barely audible. 独木舟的头部触到了湖岸的砂砾,发出了隐约可闻的沙沙声。
- A gravelly voice. 沙哑、刺耳的嗓音
- She heard a gravelly male voice. Kevin, desperate, had also called 911. Deborah asked his name and location. 是一个男人冷森森的声音。绝望中的凯文,也拨打了911。德博拉询问了他的名字和所处的位置。
- I prefer a sandy beach to a shingly one. 我喜欢沙滩,不喜欢遍布小圆石的海滩。
- The beach is a good place to fish from. 海滩是钓鱼的好地方。
- He went down to the beach for a swim. 他去海滩游泳了。
- We had a lot of fun at the picnic on the beach. 我们在海滩上野餐,趣味无穷。
- We spent a heavenly day at the beach. 我们在海滨痛痛快快地玩了一天。
- The final 20 kilometres, like some sort of supreme ordeal before we reach our goal, is tricky driving: a gravelly one-vehicle track down the mountain, with a precipitous drop on one side. 最后20公里路十分难走,如同到达目标前的某种最残酷的考验:沿着山边的一条仅容一车通过的碎石路,另一边是险峻的陡坡。
- Billy Lodi抯 mat career might not be as impressive as Cameron抯, but he is a natural: home-skilled, sexy as fuck, tough and mean, a gravelly, growling voice with nasty attitude for days. 比利洛迪抯垫的职业生涯可能不会令人印象深刻的卡梅伦抯,但他是一个自然:家庭技能,性感的他妈的,艰难,意味着,一个砾石,咆哮的声音与讨厌的态度好几天。
- We were all sunburned from a day on the beach. 我们在海滩上过了一天,皮肤都晒黑了。
- I prefer a sandy beach to a shingle one. 我喜欢沙滩,不喜欢遍布小石子的海滩。
- I saw a rotting hulk on the beach. 我在海滩上看见了一艘腐烂的废船。
- There is a vein of melancholy in his character. 他的性格中有少许忧郁的气质。
- There was a long lag in forwarding mail to us. 转寄给我们的邮件晚了很久才拿到。
- A bowl of hot chicken soup is good for you. 喝一碗热鸡汤对你有好处。
- A cloud is a mass of vapor in the sky. 云是天空中的水汽团。
- A country must have the will to repel any invader. 一个国家得有决心击退任何入侵者。
如果不是以上词条,让海词编辑来提供解释 请改进词条
- 今日热词


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The nature of online marketing is interesting, complex and always evolving. There are a great many techniques you can use that can be customized to suit your own specific business. This can make starting out a bit tricky for a novice. This list will help prepare you for this challenge.
Before you set out to get your website ranked, you should concentrate on building a quality site. This is priority number one for all webmasters, regardless of their business niche. Attention to detail is important when creating your site. The effort that you put into making your site look great and work smoothly will pay off, as you will have less work to do down the road.
Watching your competition will help you immensely in internet promotion. Whatever aspect of the market you're poised to capitalize on, others will be working that angle too.
Advertise your company on popular websites online. This may be a big investment, but it can make a great big difference. Look for a site that gets lots of traffic, and this can help you get your name out there.
Be as content focused and through as you can when you create a website. You must provide knowledge in a small amount of time so that your prospects understand what they are going to buy. Repeat information, fluff and unreliable information should be avoided.
Video marketing is a great choice for Web marketing. Use keywords in the descriptions and captions to increase the visibility of your website. Remember that you have to entice customers into watching the video in order to realize the benefit; a compelling title and an intriguing picture are important.
The first strategy worth looking into when it comes to affiliate marketing is to scrutinize your competition. Check out other websites in your field to discover what they could improve upon. If your competitors are neglecting a certain approach, take advantage of this by developing this aspect of your website.
There are so many ways you can piece together an Affiliate marketing strategy that it is both an amazing benefit and a potential detriment. It is a good thing that there is a lot of room for customization, but that means it can be very difficult to make your choices. The tips and techniques discussed in this article will help you sort through all the steps associated with Web marketing.


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you are here issue #1
$5.00 / Sold Out
Shot on 35mm black and white film, 'You are here' depicts everyday life in Los Angeles through 21 never before released images.A juxtaposition of people and places within time and situations of a major city killing itself to adapt to ever changing individuals, us.
Developed and printed in my studio, including a hand sewn organic cotton binding; hand waxed for durability. You are here issue #1 comes in an edition of five. signed and numbered.


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Archive for the 'ram 2500' Tag
2019 Ram Promaster
Furnace inspection, extra insulation, lawn and garden cleanup – these are just some of the Fall winterizing tasks in full swing around the country. Busy contractors need tools they can count on to get the job done the first time, every time. And one tool they count on the most is their truck.
Ram Commercial vans and 3500, 4500 and 5500 Chassis Cabs are built to handle big payloads, tow heavy loads and efficiently adapt to each contractor's specific needs with class-leading capability and a host of class-exclusive features. You've seen these hardworking trucks on the road all over town. Now, #TruckThursday kicks off a five-week video series focusing on the capability and features of this hardworking truck lineup, beginning with the 2019 Ram ProMaster.
Ram Trucks Launched Heavy Duty Night Models
Ram Trucks revealed both of the Heavy Duty models for the first time at the 2017 Chicago Auto Show. Based on the Sport appearance package, the Night adds blacked-out features, including wheels, grille surround and badging, to deliver an eye-catching twist on the performance-enthusiast package.
If you're looking for a way to stand out from the crowd – these Heavy Duty Night models will do just that.
The new Ram Heavy Duty Night package adds:
Black grille surround with black billet inserts
Bold "RAM" tailgate with blackened-out lettering from Ram Power Wagon®
Black Ram's head grille badge
Flat black Ram 2500 or 3500 Heavy Duty door badge
Flat black 4×4 tailgate badge (if applicable)
Flat black powertrain door badging
The Night package is available on Ram 2500 and 3500 (single-rear-wheel models only) in Crew Cab configurations, 4×2 or 4×4, and any available powertrain combination (5.7L HEMI® V8, 6.4L HEMI V8 or 6.7L Cummins® I-6).
Ram 2500 and 3500 HD Night models are available in Bright Silver Metallic, Bright White, Brilliant Black Crystal, Delmonico Red Pearl and Granite Crystal Metallic.
Production of the 2017 Ram HD Night models began early February 2017. Ram Heavy Duty Night pricing starts at $45,520 MSRP, plus $1,320 destination.
Read more at: https://blog.ramtrucks.com/features/ram-trucks-launched-heavy-duty-night-models/
Master Every Job with the 2016 Ram 2500
No matter what the workday calls for, you can rest assured that when you're behind the wheel of the 2016 Ram 2500 you can tackle it all, and then some. We at Dick Scott Chrysler Dodge Jeep Ram and Dick Scott Motor Mall know you will absolutely be impressed by all its power, refinement, and dynamic features once you take this tough pickup for a test drive.
To be a truck that is ready for any sort of job, you need to have an impressive engine, and the 2016 Ram 2500 has three available powertrains to choose from, depending on your needs. Coming standard is the 5.7L HEMI® V8 with Variable Valve Timing. When you put the pedal to the metal, you'll have 383 horsepower, 400 lb.-ft. of torque, a 13,890-lb. towing capacity1 for bringing a boat to Lake St. Clair, and a 3,060-lb. payload capacity1. You also have the choice of the 6.4L HEMI® V8 with best-in-class gasoline horsepower and torque with 410 horsepower and 429 lb.-ft. of torque. It also boasts a 16,320-lb. towing capacity1 and a 3,990-lb. payload capacity. If you prefer diesel, the 6.7L Cummins® Turbo Diesel choice holds nothing back with 370 horsepower and a whopping 800 lb.-ft. of torque. You'll also have a best-in-class 17,980-lb. tow rating in addition to a 3,160-lb. payload rating.
The 2016 Ram 2500 helps you make the most out of all this impressive power with a variety of features. For instance, the standard class-exclusive2 5-link coil rear suspension works to reduce overall friction without limiting your towing capabilities for improved turning ability and an enhanced ride.
You'll also have the trailer harness connector and heavy duty hooks for control over whatever it is you need to bring with you. For smaller items, take advantage of the available class-exclusive RamBox® Cargo Management System. This toolbox is built-in to the side rails of the truck bed, and is lit, secure, durable, and drainable. Also within the truck bed are box rail caps to reduce the number of scratches and scrapes that occur when unloading cargo, and the Ram Truck Tailgate Lift Assist to make lowering your tailgate smooth and easy.
To go with all this power and functionality is an interior you'll actually enjoy spending time in. Like the versatile exterior, 2016 Ram 2500 provides an abundance of room for a variety of cargo. For instance, the standard 40/20/40 split-folding rear bench can easily accommodate all your gear, while the built-in under-seat storage on Crew and Mega Cab® trims keep the essentials close at hand. When the winter months are upon us, take advantage of the available heated steering wheel and heated front and rear seats. Up front, you'll find the available largest-in-class 8.4-inch touchscreen display with Uconnect®. Here, you can use SiriusXM® Satellite Radio, Hands-Free Calling, and even available navigation. Also on this screen is a view from the available ParkView® Rear Back Up Camera, helping you to hook up your trailer and reverse out of a tight parking space at The Home Depot.
You are sure to be confident in your ability to tackle any task with the 2016 Ram 2500. To see all its impressive power and tough features, visit Dick Scott Chrysler Dodge Jeep Ram or Dick Scott Motor Mall to take one for a test drive. Our new-vehicle showroom is open six days a week, so it's easy to find time to check out this commanding truck.
2017 Ram Power Wagon Laramie caught
Brian Williams caught this 2017 2500 Ram Power Wagon Laramie while it was being tested in high altitudes in the Rockies. He wrote that it's only missing the legacy "Macho Power Wagon" graphics and blacked out bits.
The brawny, off-road-oriented Power Wagon, revamped for 2017, will gain a more luxurious version for those who need the muscle but also want the pampering of the wood and leather-lined high-end trim offered on other Ram pick-ups.
The Laramie will get more brightwork as befitting of this top-spec Ram, with a new take on the Power Wagon's Rebel-derived grille – a sort-of chromed mesh insert replaces black plastic. Chromed wheels are now wrapped in the chunky off-road tires, too.
Inside, the Power Wagon should get the typical Laramie treatment, which means upgraded leather with thick stitching and wood-like trim.
Expect to see the Laramie Power Wagon later this year.
Read more at: http://www.allpar.com/news/2016/06/2017-ram-power-wagon-laramie-caught-31987
Changes for the 2016 Ram HD pickups
A reliable source oh2o provided a list of what will be changing for the 2016 model year in the Ram 2500 and Ram 3500 pickup trucks.
Production of the 2016 model year pickups begins on July 20, 2015, and will include the new Limited, with the same controversial grille and tailgate appearance as the Ram 1500 Limited (shown below). Available as a crew cab or mega cab only, it will have unique wheels (except on double-rear-wheel setups, or "duallies," which will carry over the old wheels). The interior is all-black and similar to that of Ram 1500 Laramie Limited; bumpers are painted, with optional chrome.
The photo above shows a 2016 Ram 2500 Laramie Limited, as shown at the Chicago Auto Show. Other models are likely to have a grille similar to those of the 2015 trucks.
All 2016 "heavy duty" Ram pickup trucks will have a new center console layout; Longhorn and Limited will also use a new, wooden tambour (sliding console) door.
New 18 inch aluminum wheels will be used on Big Horn, Lone Star, and Outdoorsman, with new pickup box lighting (LED lights on each side at the rear) on Longhorn and Limited, optional on some other models, and included with Luxury Group and Rambox.
Engines appear to remain the same, except for a new compressed natural gas (CNG) option on Crew Cab / long-bed in rear wheel drive, and regular cab models in rear and four wheel drive. Ram continues to have the only factory-upfitted CNG option, not using an aftermarket solution.
The base radio on Tradesman remains the same but will be renamed from RA1 to Radio 3.0 AM/FM. Some models will get optional front and rear parking assistance (standard on Laramie and higher trimlines).
phablet holderThe rear camera will be able to toggle between a cargo view and the backup camera; the cargo view camera will be packaged in the Convenience Group with rain-sensing wipers and automatic-beam headlamps.
Finally, the usual paint changes are going to be made, with red pearl replacing deep cherry red, "Luxury Brown" replacing "Western brown," and a new pearl white option on higher-level trucks starting with September production.
It normally takes a month or two for trucks to reach dealers after production begins.
As read on: http://allparnews.com/index.php/2015/05/changes-for-the-2016-ram-hd-pickups-28681
Ram grille change no big deal?
Opinion/Analysis: it seems to me that the uproar over the new grilles on the Ram Laramie and Rebel is a classic case of making a mountain out of a molehill: a task made more difficult because it starts without a molehill.
Automotive News' Larry Vellequette and Allpar's Daniel Bennet have written about the change, saying it's a sign that the Ram brand is suffering from an identity crisis. Their argument is that Ram trying to further distance itself from Dodge by moving away from the crosshair grille and the ram's head badge.
The more likely reason for the change is to let buyers know this is a new Ram pickup [as alluded to by designer Greg Howell].
How long has the Ram pickup used the same general grille design with only very small changes that most consumers won't even notice? Since Ram became a separate brand five years ago? In fact, the same general grille design was used on the last model year of the Dodge Ram pickup.
Maybe it's time for a change?
The fact the new grille's first appearance on a regular production truck came on the top-of-the-line Laramie is an indication that Ram is looking to persuade owners of earlier premium Ram pickups to trade their old truck by making the new truck visibly different.
When I was young, grille changes were an annual event, making each new model distinct from those that came before. Automakers don't do that any more, but a change after six years seems reasonable.
The use of the prominent "RAM" on the Rebel's grille is similar to what Ford has done on the Raptor, which has a big "FORD" on the grille instead of a blue oval. It sets the special truck apart from other models.
As for the large RAM on the tailgate, name a pickup brand that hasn't done this at one time or another.
In short, there's no identity crisis required to explain the change.
As far as establishing a brand identity, I would imagine that if you asked most male consumers to complete the phrase "Guts, Glory…", most would say "Ram." They might even try to sound like Sam Elliott. That's successful branding.
Worries about losing the classic Ram logo would seem to be unfounded. A look at the interior shows the familiar shield is right in the center of the steering wheel. Considering that it's much less costly to change a small badge than it is to change a grille and tailgate, one would assume any effort to rebrand would include that change.
Consider the ProMaster City. While the big ProMaster was already in production, it would have been easy to change the small van's grille and badging. Yet the ProMaster City has Ram shield badges front and rear.
Now consider at the two brands' product lines: Ram has pickups, chassis-cabs and commercial vans (ProMaster City is clearly targeted at businesses). Dodge has passenger cars, family minivans and SUVs. The only Dodge fleet vehicles are special purpose, such as the Charger Pursuit and the Durango SSV. There isn't any overlap, even in the same showroom.
It was Chrysler, not Fiat, that originally pushed for a retail network in which as many dealers as possible sold all the Chrysler brands.
If anyone is worried that Ram is phasing out the Ram logo, the first question that comes to mind is "Why would they?" The brand name is Ram; what else are they going to use?
It's unlikely that Bob Hegbloom, Sergio Marchionne or Olivier Francois lose much sleep over whether dealers or consumers call the truck a Ram or a Dodge Ram. FCA US and people can call it anything they want as long as it changes hands from the first group to the second group in large quantities, and Ram U.S. sales last year were the best since the all-time record year of 2003 and missed setting a new all-time Graham/Dodge/Ram sales record by just 9,583 sales.
In the end, the grille change isn't a quest for identity or an escape from the shadow of another brand. It is a relatively inexpensive styling change made by a brand that seems comfortable enough in its own skin to try something new.
Read more at: http://www.allpar.com/news/index.php/2015/03/relax-ram-grille-change-no-big-deal
Black is the new black
Some Web sages noticed that the only interior color available for the Ram Rebel and Laramie Limited was black (this has been true for both generations of Laramie Limited). With the 2015 Ram 1500 Laramie Limited, even the headliner and pillars are black.
We asked Ram lead interior designer whether this was due to customer choice or cost, and he said it was partly a matter of being appropriate to the vehicle.
There's just something about wearing a black suit, just a nice black suit, that your details pop. I think that's the thing that we've learned over the years, that it's okay to have a very calming black, very consistent, and then accent it with colors. That's what we've chosen for like you mentioned the Limited; we've chosen it for the Rebel that we're sitting in. It's appropriate.
Mr. Nagode also mentioned the dirt resistance (to fingerprints) in truck use, as well as customer choice if presented with different options at a lot. However, he also noted that for other vehicles, different colors were more appropriate, particularly in cars such as Charger, Challenger, and Viper. The Laramie Longhorn, he pointed out, comes in either black with brown seats, or in "a frost color" with a warmer brown.
It's just sometimes the extremes, we tend to kind of limit the choices. Like our Express and Tradesman really only comes in a black environment. And again, that's perfect for someone that's going to get it dirty. … At the core of our market, we tend to offer more colors, more variations.
As read on: http://www.allpar.com/news/index.php/2015/02/black-is-the-new-black
Ram Jam: Ram Trucks CEO Talks Hellcat, Compact Trucks, and More
Last August, longtime Chrysler executive (he's been with the company since 1988) Robert Hegbloom took the position of CEO of Ram Trucks. Now that he's had time to get comfortable with the reins, we sat down to talk sales numbers, EcoDiesel, mid-size trucks, and more—much more.
Car and Driver: Ram sold 395,567 trucks in 2014, placing it in fourth place in the bestselling-vehicle derby behind the Toyota Camry (396,988), the Chevrolet Silverado (471,918), and the Ford F-series (679,496). Safe to call it a big year?
Robert Hegbloom: We were up 28 percent year over year, and three points overall in the market. Basically, we look at our customers and see that they play in a number of areas: work, outdoor [recreation], motorsports, and even [just using the truck] as a family vehicle. And in those spaces, customers are looking for something unique. But the first thing with a light-duty truck is fuel economy, which is so different from where it was a few years ago. We want to deliver on fuel economy first, and then, depending on what their particular needs are, make sure we have the right package.
Might the product lineup get diluted by the staggering amount of available trims and packages and submodels? Do you see consumers being overwhelmed by the number of choices?
You know, it varies when regionalism comes into play. For instance, when you go down to Texas, the Laramie Longhorn is very popular, and there are some urban areas like Dallas, Fort Worth, Houston, and Austin, too, and they like chrome. Now when you go to California, they like the monochromatic look. You go to another part of the country and they have preferences, too. So you'll have regional differences among the package preferences, and the dealers aren't going to stock all of them.
We know this is sensitive subject, but now that the F-150 has come out and the numbers have posted, where is Ram on the use of aluminum? Last time we talked, [Ram vehicle line executive] Mike Cairns said "aluminum is for beer cans."
Well, we are delivering fuel economy using technology, like the EcoDiesel and the Pentastar V-6 mated to the eight-speed transmission. Then you add the enablers we put in place such as active grille shutters and aerodynamics. We still have the best aero in the segment, and we want to stay consistent there while looking for new opportunities in efficiency.
Are the guys who buy diesel Ram trucks concerned with being green, or is it just about torque and fuel efficiency?
Ultimately, it's about delivering on fuel economy without sacrificing any capability. You get 240 horsepower and 420 lb-ft of torque and 29 mpg on the highway [with the EcoDiesel]. And to put that in perspective, consider that when we launched the 5.9-liter Cummins [in 1989], it had twice the displacement, yet made less horsepower and less torque. The 3.0-liter diesel delivers all the capability a light-duty buyer demands. Typically, consumers see a label and think they aren't going to achieve the quoted mileage; what we're seeing is customers writing [to us] about their EcoDiesel truck and saying they are getting 29 mpg or even better.
Gas prices are currently at the lowest they've been in years. Is that slowing the sales of the EcoDiesel?
Back in December, when gas prices were at their lowest, we had the highest-selling month YTD that we've had since the introduction of the EcoDiesel. We are at 26 days on lot with the EcoDiesel, even being a full year into the model's availability and while we are increasing our volume. Typically we see much more fluctuation with gasoline prices than with diesel prices, so it doesn't affect sales as much.
Is there a cap on the number of EcoDiesels you can produce?
We are trying to add 20 percent to our production. We received enough orders on this thing right out of the gate that took us almost to the model year [to have enough available units for sale]. You expect to get a big lift when you come out with something like that, but to sustain it for a year, that's really something.
Now the big question: Hellcat Ram?
You know, we did an SRT a number of years ago, and it was something that at the time worked for a little bit. We get a few people who think a Hellcat truck would be fun, we could take it to Woodward Avenue and have some fun, but there's not really a big appetite for that type of performance vehicle right now. Plus [Dodge chief] Tim Kuniskis wants all the Hellcat motors he can get.
ProMaster City—what's the early feedback?
We need to get them out there. We started shipping in the end of December, and the people that have driven them have been pleased. The 2.4-liter Tigershark four-cylinder with the nine-speed transmission surprised a lot of people, as did the independent suspension, which we were very focused on, because we wanted the driving characteristics right. All three of those are major changes compared to what is in Europe. We took a great platform and optimized it for our market. I'm pretty excited to get them here.
Is a diesel option for the ProMaster City in the cards?
They have it in Europe, and the first time I ever drove one it was a diesel with the manual. But time will tell. We have the technology, but you start getting into the cost of ownership, which is a major driver in a commercial-oriented package. Acquisition price plays a big role in that, and I'm just not sure the appetite is there for that.
What's the possibility that a mini/mid-size truck based on the Fiat Strada will make it to the U.S.?
No plans. It's a nice package, but to meet all the homologation requirements, you would essentially have to start over. We looked at it, it's a 200,000–300,000 unit a year segment with a number of players in it, so unless you see some major changes there's just not a big opportunity. [A new generation is due next year. Hmm . . . —Ed.]
As read on: http://blog.caranddriver.com/ram-jam-ram-trucks-ceo-talks-hellcat-compact-trucks-and-more/
2015 Ram 1500 Review
The 2015 Ram 1500 ranks 1 out of 5 Full Size Pickup Trucks. This ranking is based on our analysis of published reviews and test drives of the Ram 1500, as well as reliability and safety data.
The 2015 Ram 1500 impresses reviewers with its best-in-class ride, excellent cabin materials, responsive transmission and class-exclusive diesel engine option.
Because it has the best combination of positive reviews and long-term ownership costs in its class, the Ram 1500 is the 2015 U.S. News Best Full Size Truck for the Money.
The Ram 1500 has a base V6 that auto critics say has sufficient power. The optional V8 and turbodiesel V6 both earn praise for their ample power, and reviewers say either engine is a good choice for towing and hauling heavy loads. An eight-speed automatic transmission is standard, which reviewers praise for its responsiveness and smooth shifts. Fuel economy for a base Ram 1500 is an EPA-estimated 17/25 mpg city/highway, which is comparable with rivals' estimates. The diesel model returns 20/28 mpg, which is excellent for the class. Test drivers say the 2015 Ram 1500 sets the standard for ride comfort in the segment, and they add that its optional air-adjustable suspension is helpful for keeping the truck level when towing or hauling heavy loads.
Auto journalists are impressed with the soft-touch materials and build quality of the 2015 Ram 1500's cabin. They say the interior is quiet at speed and note that the front seats are comfortable and supportive. In either Quad or Crew Cab configuration, both of which seat up to six, reviewers report that there is plenty of legroom. The Ram 1500 offers an abundance of storage spaces in the cabin, reviewers write, and they like its available lockable RamBox storage compartments for storing smaller cargo in sides of the bed. A USB port and auxiliary input jack are standard. Optional features include navigation, satellite radio, a rearview camera and a Uconnect infotainment system with Bluetooth and voice control capability. Test drivers write that Uconnect is very user-friendly and they appreciate that there are redundant physical buttons for most major climate and audio adjustments.
"The Ram 1500 also offers a wide choice of trim levels, from workhorse, to street cruiser, to luxury liner. Even if you're a die-hard loyalist to another brand, you still owe it to yourself to check out these outstanding trucks." — Consumer Guide
"Available with all the required cab and bed configurations, in either 2-wheel or 4-wheel drive, the Ram 1500 goes above and beyond with its stylish exterior and well-appointed cabin." — Kelley Blue Book
"The Ram 1500 has the nicest interior of any full-size pickup. Its upgraded touchscreen interface is impressively easy to use and offers substantial technology capabilities. We're also fond of the Ram as it offers a composed and smooth ride whether you're driving it on- or off-road." — Edmunds (2014)
"Already gifted with capable handling and handsome interiors, the Ram 1500 lineup is now even more unique and appealing with the addition of the EcoDiesel option. It isn't the motor for everyone – the cheaper, punchier and more characterful Hemi is still a stellar choice – but its combination of low-end torque and efficiency will undoubtedly win over many hearts and wallets. We expect Ram will have little problem achieving its goal of a 15 percent take rate for the EcoDiesel." — Left Lane News (2014)
Read more at: http://usnews.rankingsandreviews.com/cars-trucks/Ram_1500/


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Amazonas Florestal Ltd. Announces Its Wholly-Owned Subsidiary, Amazon Hemp Ltd., Achieves Registration With The Colorado Department of Agriculture; Moves Closer to Meeting Requirements for Planting of 100 Acre Plot with Industrial High CBD Hemp7:21 AM March 15, 2017
DENVER, CO–AZFL, (Marketwired – Mar 15, 2017) – Amazonas Florestal, Ltd. (www.azflusa.com) (OTC PINK: AZFL), a natural resources company dedicated to innovative, sustainable forest management, Industrial CBD Hemp and the certification and sales of carbon credits today announced its wholly-owned subsidiary, Amazon Hemp Ltd, has moved one step closer to meeting requirements for planting of its 100 Acre Plot with its Industrial High CBD Hemp by achieving registration with the Dept. of Agriculture for the State of Colorado. The company is closeRead more