Alejandro ; Montecinos

Alejandro E Montecinos

Lucia Couceiro

Akira F Peters

Antoine Desrut

Myriam Valero

Marie-Laure Guillemin
email: marielaure.guillemin@gmail.com

SPECIES DELIMITATION AND PHYLOGEOGRAPHIC ANALYSES IN THE ECTOCARPUS SUBGROUP SILICULOSI (ECTOCARPALES, PHAEOPHYCEAE) (1)

Keywords: barcode, brown alga, COI-5P, cryptic species, <i>Ectocarpus</i>, hybridization, introgression, ITS1, species delimitation barcode, brown alga, COI-5P, cryptic species, Ectocarpus, hybridization, introgression, ITS1, species delimitation ABGD, Automatic Barcode Gap Discovery BI, Bayesian Inference COI-5P, 5

The genus Ectocarpus (Ectocarpales, Phaeophyceae) contains filamentous algae widely distributed in marine and estuarine habitats of temperate regions in both hemispheres.

While E. siliculosus has become a model organism for genomics and genetics of the brown macroalgae, accurate species delineation, distribution patterns and diversity for the genus Ectocarpus remain problematic. In this study, we used three independent species delimitation approaches to generate a robust species hypothesis for 729 Ectocarpus specimens collected mainly along the European and Chilean coasts. These approaches comprised phylogenetic reconstructions and two bioinformatics tools developed to objectively define species boundaries (GMYC; General Mixed Yule Coalescence Method and ABGD; Automatic Barcode Gap Discovery). Our analyses were based on DNA sequences of two loci: the mitochondrial cytochrome oxidase subunit 1 (COI-5P) and the nuclear internal transcribed spacer 1 of the ribosomal DNA (ITS1). Our analyses showed the presence of at least 15 cryptic species and suggest the existence of incomplete lineage sorting or introgression between five of them. These results suggested the possible existence of different levels of reproductive barriers within this species complex. We also detected differences among species in their phylogeographic patterns, range and depth distributions which may suggest different biogeographic histories (e.g., endemic species or recent introductions).

Introduction

Delineating species boundaries is a longstanding methodological and conceptual challenge, especially in algal systems. Some of the problems arise from the fact that species are dynamic entities that change with time (Sites andMarshall 2003, 2004) and a plethora of species concepts has been proposed [START_REF] Mayden | A hierarchy of species concepts: the denouement in the saga of the species problem[END_REF][START_REF] Mayden | Consilience and a hierarchy of species concepts: advances toward closure on the species puzzle[END_REF][START_REF] Coyne | Speciation[END_REF]. Recently, a new line of thinking was put forth among biologists whereby species are considered as scientific hypotheses (species hypotheses) and species delineation is a process of refutation based on the acquisition of new evidence [START_REF] Pante | Species are hypotheses: avoid connectivity assessments based on pillars of sand[END_REF].

DNA-based methods, such as single gene barcoding, have been proven especially useful to uncover cryptic species where classical taxonomy has been problematic in organisms characterized by simple morphology and / or by high phenotypic plasticity (e.g., in animals: Yamashita and Roads 2013; in plants: Carstens and Satler 2013; in seaweeds: [START_REF] Tronholm | Species delimitation, taxonomy, and biogeography of Dictyota in Europe (Dictyotales, Phaeophyceae)[END_REF]. However, problems are linked with this single gene approach.

Indeed, single-locus data represent the history of a single gene that might not be representative of organismal history. Differences in the overall amount of differentiation between loci or in how these loci reconstruct the relationships among groups generate discordance in species boundaries delineation among markers and the importance of using multiple independent loci to generate robust species hypotheses has been repeatedly emphasized [START_REF] Dupuis | Multi-locus species delimitation in closely related animals and fungi: one marker is not enough[END_REF].

Distance-based approaches have classically defined species using arbitrary thresholds (universal or defined visually using a barcode gap in a particular group of species, Hebert et al. 2003). Species have also been defined based on the existence of wellsupported monophyletic groups [START_REF] Wiens | Delimiting species limits in spiny lizards (Sceloporus)[END_REF]. However, monophyly, while a discrete criterion, is arbitrary with respect to taxonomic level [START_REF] Goldstein | Integrating DNA barcode data and taxonomic practice: determination, discovery, and description[END_REF]. Methods characterized by an increased statistical rigor and better objectivity in delimiting species, such as the General Mixed Yule Coalescent (GMYC) (Pons et al. 99 2006, Monaghan et al. 2009) and the Automatic Barcode Gap Detection (ABGD) (Puillandre et al. 2012a), were recently developed to detect discontinuities in DNA sequence variation associated with species boundaries. GMYC uses a pre-existing phylogenetic tree to determine the transition signal from speciation to coalescent branching patterns. While, ABGD detects the breaks in the distribution of genetic pairwise distances, referred to as the 'barcode gap', relying exclusively on genetic distance between DNA sequences. GMYC and ABGD analyses combined with searches for well-supported monophyletic groups in phylogenetic reconstructions have been used to detect the existence of cryptic species in many taxa (e.g. snails: [START_REF] Prevot | Exploring species level taxonomy and species delimitation methods in the facultatively self-fertilizing land snail genus Rumina (Gastropoda: Pulmonata)[END_REF]fish: Alò et al. 2013;copepods: Cornils and Held 2014;red algae: Payo et al. 2012 and[START_REF] Pardo | A multilocus species delimitation reveals a striking number of species of coralline algae forming maerl in the OSPAR maritime area[END_REF] or brown algae: [START_REF] Vieira | Toward an inordinate fondness for stars, beetles and Lobophora? Species diversity of the genus Lobophora (Dictyotales, Phaeophyceae) in New Caledonia[END_REF].

The genus Ectocarpus Lyngbye (Ectocarpales, Phaeophyceae) is widely distributed in marine and estuarine habitats of temperate regions in both hemispheres [START_REF] Stache | Sexual compatability and species concept in Ectocarpus siliculosus (Ectocarpales, Phaeophyceae) from Italy, North Carolina, Chile, and New Zealand[END_REF].

Ectocarpus spp. is found as a short-lived annual and often colonizes abiotic substrata or grows as an epiphyte on macrophytes; the habitat of Ectocarpus spp. includes the subtidal up to high intertidal pools (Russell 1967a(Russell , 1967b(Russell , 1983a(Russell , 1983b)). Members of Ectocarpus spp. complex have been described as important contributors to biofouling and are frequently encountered as epiphytes in mariculture settings [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF]). The genus Ectocarpus has a long research history, starting in the XIX century with the first taxonomic descriptions of this genus [START_REF] Dillwyn | British Confervae; or colored figures and descriptions of the British plants referred by botanists to the genus Conferva[END_REF][START_REF] Lyngbye | Tentamen hydrophytologiae danicae; continens omnia hydrophyta cryptogama Daniae, Holsatiae, Faeroae, Islandiae, Groenlandiae hucusqve cognita, systematice disposita, descripta et iconibus illustrata, adjectis simul speciebus norvegicis. Opus, praemio ab Universitate regia hafniensi ornatum, et sumtu regio editum. Hafniae, typis Schultzianis[END_REF].

Despite being a model organism [START_REF] Peters | Proposal of Ectocarpus siliculosus (Ectocarpales, Phaeophyceae) as a model organism for brown algal genetics and genomics[END_REF], Cock et al. 2010), basic knowledge concerning species delineation, distribution patterns, diversity and differentiation remains elusive (Peters et al. 2010a).

6

Initial morphology-based descriptions of species diversity have a long and controversial history. For example, Hamel (1931Hamel ( -1939) ) recognized five species along the European Atlantic coast, which he classified into two major groups based on branching pattern and sporangium shape: the section "siliculosi" and the section "fasciculati". Later, [START_REF] Cardinal | Étude sur les Ectocarpacées de la Manche[END_REF], using field material from the French Channel, proposed another classification and distinguished four species with seven varieties in the Ectocarpus subgroup siliculosi and three varieties in the Ectocarpus subgroup fasciculati.

Conversely, [START_REF] Russell | The genus Ectocarpus in Britain. I. The attached forms[END_REF]Russell ( , 1967a) ) using isolates from around the British Isles, demonstrated that sporangium morphology was not an informative species character.

His proposal to reduce the number of European species to two (E. fasciculatus and E. siliculosus) was later supported by crossing and chemical studies [START_REF] Müller | Crossing experiments, lipid composition, and the species concept in Ectocarpus siliculosus and E. fasciculatus (Phaeophyceae, Ectocarpales)[END_REF] and was the most widely accepted classification system until recently (but see Peters et al. 2010a). Nonetheless, within E. siliculosus, reproductive barriers have been reported between isolates from different geographical areas (reviewed in Stache-Crain et al. 1967). Prezygotic barriers have been described for populations from NE America [START_REF] Müller | Sexual isolation between Europe and an American population of Ectocarpus siliculosus (Phaeophyta)[END_REF]; likewise, reduced development or normal sporophyte development with inhibition of meiosis (post-zygotic barriers) has been observed for strains isolated from different hemispheres [START_REF] Müller | Sexual reproduction in British Ectocarpus siliculosus (Phaeophyta)[END_REF][START_REF] Müller | Genetic affinity of Ectocarpus siliculosus (Dillw.) Lyngb. from the Mediterranean, North Atlantic and Australia[END_REF][START_REF] Müller | Studies on sexual compatibility between Ectocarpus siliculosus (Phaeophyceae) from Chile and the Mediterranean Sea[END_REF][START_REF] Stache | Sexual compatability and species concept in Ectocarpus siliculosus (Ectocarpales, Phaeophyceae) from Italy, North Carolina, Chile, and New Zealand[END_REF]). Despite these observations, [START_REF] Müller | Sexual reproduction of Ectocarpus siliculosus (Ectocarpales, Phaebphyceae) in Japan[END_REF] proposed to collapse E. siliculosus isolates into a single species arguing that full or slightly reduced interbreeding patterns could be explained by the geographical isolation between populations in this world-wide distributed species. However, this explanation has been questioned by different studies where sequence-based analyses have identified cryptic diversity within the Ectocarpus genus. First, phylogenetic analyses using ITS1 (nrDNA) and the Rubisco spacer of choloroplast DNA (cpDNA) of 43 Ectocarpus strains isolated 7 from all continents except Antarctica, showed several lineages within the Ectocarpus subgroup siliculosi [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF]). Second, using three additional markers (cox3 and rps14-atp8 (both from mtDNA) and ITS2 (nrDNA)) and including samples isolated from NW France, Peters et al. (2010a) suggested the existence of at least four different lineages within the Ectocarpus subgroup siliculosi; for one of the four lineages, they proposed to reinstate the name E. crouaniorum Thuret coined by Thuret in Le Jolis. Third, the presence of most lineages described by [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF] was later confirmed for strains sampled along the South-East Pacific coast (Peters et al. 2010b). Finally, using a single-locus approach (COI-5P) on samples from NW France, Mediterranean Sea and Asia, [START_REF] Peters | Barcoding of cryptic stages of marine brown algae isolated from incubated substratum reveals high diversity in Acinetosporaceae (Ectocarpales, Phaeophyceae)[END_REF] reported again several lineages previously described in [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF] as well as 14 additional lineages possibly representing different species. However, this last study also warned against problems linked to a single-gene approach, such as incomplete lineage sorting or introgression.

The molecular findings discussed above support the probable occurrence of highly divergent genetic lineages, including cryptic species, within the Ectocarpus section "siliculosi". However, none of the previous studies has employed an integrative approach to clarify the species diversity within Ectocarpus and evaluate introgression levels within and among natural populations. The present study uses two unlinked loci (i.e. COI-5P and ITS1 DNA-markers) and a set of methods developed to delimit species to clarify the number of cryptic species within this group using 729 specimens collected mainly along the European and Chilean coasts. The extent to which natural hybridization and introgression occur in the field was investigated by searching for incongruence between the independent nuclear and mitochondrial markers. Finally, 8 phylogeographic patterns, range and depth distributions of the most common Ectocarpus species were studied.

Materials and Methods

Field collections and isolation of Ectocarpus strains

Seven hundred and twenty-one Ectocarpus samples were collected from 37 sites located along the North-East Atlantic (NEA), Mediterranean and South-East Pacific (SEP) coasts. They were complemented with 8 strains isolated from USA, South Korea, Australia and New Zealand (Table 1). Samples collected in the field were isolated and maintained as clonal cultures, as described in [START_REF] Couceiro | Evolution and maintenance of haploid-diploid life cycles in natural populations: the case of the marine brown alga Ectocarpus[END_REF]. Position on the shore of the collected individuals was recorded using a coarse classification (high intertidal, H; mid intertidal, M; low intertidal, L; upper subtidal, US; subtidal, S; or drifting, Drift), in order to examine whether putative cryptic species occupy different tidal zones.

DNA extraction, sequencing and alignments

Total DNA was extracted from lyophilized samples using the NucleoSpinR 96 Plant Kit (Macherey-Nagel, Duren, Germany). Partial COI (COI-5P, mitochondrial) was amplified using the primers GAZF2 and GAZR2 [START_REF] Lane | A molecular assessment of northeast Pacific Alaria species (Laminariales, Phaeophyceae) with reference to the utility of DNA barcoding[END_REF]) as described in [START_REF] Peters | Barcoding of cryptic stages of marine brown algae isolated from incubated substratum reveals high diversity in Acinetosporaceae (Ectocarpales, Phaeophyceae)[END_REF]. A nuclear fragment containing the ITS1 region and 224 bp of the flanking genes 18S and 5.8S was amplified using the primers and PCR conditions described by Peters et al. (2010a). PCR amplicons for both markers were sequenced at Genoscope facilities (Evry, France) or at Eurofinis Genomics (Ebersberg, Germany).

Individuals showing phylogenetic incongruences between markers were sequenced twice in order to discard contamination errors in the preparation of the samples before the sequencing. Sequences were aligned manually using MEGA v6.06 [START_REF] Tamura | MEGA6: Molecular Evolutionary Genetics Analysis version 6.0[END_REF]) and checked by eye; only traces with high quality values and no ambiguities were retained for further analyses.

Species-delimitation procedure

First, 729 Ectocarpus COI-5P sequences were used to define putative species within the Ectocarpus section "siliculosi" group. Among them, 710 sequences were generated in this study and deposited in GENBANK (Table 1, S1) while 19 were published by [START_REF] Peters | Barcoding of cryptic stages of marine brown algae isolated from incubated substratum reveals high diversity in Acinetosporaceae (Ectocarpales, Phaeophyceae)[END_REF] and downloaded from the same public database (Table S1). To establish putative species, two species delineation methods (ABGD and GMYC) were combined with two phylogenetic inference methods (Maximum Likelihood, ML and Bayesian inference, BI). Thereafter, putative species delineated with the COI-5P were consolidated using 630 sequences of the nrDNA marker ITS1. Five hundred and eighty sequences were generated in this study (Table 1, S2) and 50 were downloaded from GENBANK (Table S2). A single alignment including all ITS1 sequences could not be generated due to the partly high sequence variability including the presence of indels [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF]; the ITS1 dataset was therefore divided into four subgroups (see results on species consolidation below for more information about the composition of these four subgroups). Sequence alignment, tree reconstructions and ABGD tests were carried out independently for each subgroup. Sample groups were considered as species when all (or nearly all) the methods employed to test their boundaries and the results obtained for two independent genes were concordant. 10 Phylogenetic analyses were conducted separately for the COI-5P and ITS1 regions using both ML and BI methods. ML analyses were performed using RAxML v8 [START_REF] Stamatakis | RAxML version 8: a tool for phylogenetic analysis and postanalysis of large phylogenies[END_REF]. We selected the best-fit substitution model using the Akaike Information Criterion (AIC) implemented in jModelTest v2.1.8 [START_REF] Darriba | jModelTest 2: more models, new heuristics and parallel computing[END_REF]).

The selected model was GTR I+G for COI-5P, GTR +G for the first, second and fourth ITS1-sub-groups and GTR I+G for the third ITS1-sub-group. Statistical support was estimated using 1,000 replicates and a rapid bootstrap heuristic [START_REF] Stamatakis | A fast bootstrapping algorithm for the RAxML web-servers[END_REF]. BI analyses were conducted using MrBayes v3.2.3 [START_REF] Huelsenbeck | MRBAYES: Bayesian inference of phylogeny[END_REF].

Two independent analyses were run using four chains each and 20 million generations.

Trees and parameters were sampled every 1,000 generations and the default parameters for temperature and branch swapping were used. The first 20% of sampled trees were discarded as "burn-in" to ensure stabilization. The remaining trees were used to compute a consensus topology and posterior probability values. The split frequency (variance among the four independent runs) was below 0.003, confirming that the posterior probability distribution was accurately sampled.

Automatic Barcode Gap Discovery (ABGD)

ABGD identifies a limit between the frequency distribution of intra-and interspecific pairwise genetic distances, even if they overlap, by using several a priori thresholds of genetic distances chosen by the user (Puillandre et al. 2012a). Then, it is recursively applied to previously obtained groups to get finer partitions until there is no further partitioning. ABGD was remotely run at http://wwwabi.snv.jussieu.fr/public/abgd/abgdweb.html. We computed Kimura twoparameter (K2P) genetic distances among specimens using default settings.

General Mixed Yule Coalescent (GMYC)

GMYC identifies a threshold value for the shift in branching rate from coalescent lineage branching to interspecific diversification on an ultrametric tree and explicitly delimits "independently evolving" clusters (i.e. putative species) [START_REF] Pons | Sequence-based species delimitation for the DNA taxonomy of undescribed insects[END_REF], Monaghan et al. 2009). Before the analysis, duplicated haplotypes were removed from our data set using DnaSP v5.10.1 [START_REF] Librado | DnaSP v5: a software for comprehensive analysis of DNA polymorphism data[END_REF]. Branch lengths were estimated under a relaxed log-normal clock using the Bayesian analysis implemented in BEAST v1.8.2 [START_REF] Drummond | Bayesian phylogenetics with BEAUti and the BEAST 1.7[END_REF]. A coalescent (constant size) prior was used and Markov Chains Monte Carlo (MCMC) were run for 20 million generations. Trees were sampled each 1,000 generations with a 10% burn-in. A visual inspection of MCMC progression using Tracer v1.6 was performed to corroborate stabilization. An ultrametric tree was constructed using TreeAnnotator v1.8.1 [START_REF] Rambaut | TreeAnnotator version 1.6.1[END_REF]. Both the single-threshold [START_REF] Pons | Sequence-based species delimitation for the DNA taxonomy of undescribed insects[END_REF]) and the multiple-threshold (Monaghan et al. 2009) versions of GMYC were fitted on the ultrametric tree using the SPLITS v1.0-19 package for R (https://r-forge.r-project.org/projects/splits/).

Network reconstructions

Haplotype networks were reconstructed for the eight Ectocarpus species for which more than 15 sequences were available for each marker under study. The haplotype networks were reconstructed using the median-joining algorithm implemented in NETWORK v6.13 [START_REF] Bandelt | Median-joining networks for inferring intraspecific phylogenies[END_REF].

Genetic diversity

Genetic diversity indices were calculated for the three species, E. siliculosus, E. crouaniorum and Ectocarpus 6, that were sampled most frequently (i.e. >45 individuals for which both the COI-5P and ITS1 markers were sequenced). The number of haplotypes (nH); the number of polymorphic sites (S); gene diversity (H) and nucleotide diversity (π, [START_REF] Nei | Mathematical model for studying genetic variation in terms of restriction endonucleases[END_REF]Li 1979) were computed using ARLEQUIN v3.5.1.3 (Excoffier and[START_REF] Excoffier | Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and Windows[END_REF].

AMOVA analysis

For the two most widely geographically distributed species (i.e. E. siliculosus and E. crouaniorum), a nested Analysis of Molecular Variance (AMOVA, [START_REF] Excoffier | Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data[END_REF]) was implemented using ARLEQUIN v3.5.1.3 [START_REF] Excoffier | Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and Windows[END_REF] to test for the partition of genetic variance within locations, among locations within regions and among regions. Based on the geographic clustering of the sampled localities, four regions were defined: United Kingdom, France, NW Iberian Peninsula and Chile.

Results

Ectocarpus putative species delineation based on COI-5P

The 729 COI-5P sequences (603 bp) from the Ectocarpus specimens included 90 unique haplotypes with 123 variable sites and a high level of haplotypic diversity (0.9093). The phylogenies inferred from these haplotypes using ML and BI (Figure 1a) gave comparable topologies and suggested the presence of 15 putative species within the group E. siliculosi. Eleven of these lineages were highly supported monophyletic groups (bootstrap values > 84 for ML and > 0.98 for BI, Figure 1a) while the remaining four were singletons. Genetic pairwise K2P distances ranged from 0 to 0.089 and the shape of the distribution was clearly bimodal with two conspicuous maxima at 0.0025 and 0.0550. ABGD located the barcode gap within the 0.011-0.037 distance range 13 (Figure S1a) and primary partitions using this threshold suggested the existence of 15 genetic groups (Figure 1a, Figure S1b). The likelihood of the GMYC model, for both the single-and the multiple-threshold models (LGMYCsingle = 720.08 and

LGMYCmultiple = 723.40), was significantly higher than the likelihood of the null model (L0 = 704.01). However, the partitions obtained were not identical for the different threshold limits; 16 groups were delimited with the single-threshold method (confidence limits, 15-18) and 22 groups were delimited with the multiple-threshold method (confidence limits, 18-26). The likelihood values of the single and multiple threshold analyses were not significantly different, suggesting that application of the more complex multiple-threshold analysis did not result in a significant improvement of the results. Thus, we selected the single threshold over the multiple threshold model to delimit putative species in our data set (Figure 1a, Figure S2).

The eleven monophyletic groups recovered by tree reconstructions were supported as putative species by the ABGD results while only ten out of these eleven clades were supported as putative species by the GMYC single threshold results. This discrepancy involved the split of two haplotypes (L211 and L206) from the rest of the E. siliculosus clade in the GMYC (Figure 1a, Figure S2). The four singletons observed in both phylogenetic reconstructions were defined as species by both ABGD and GMYC (Figure 1a, Figure S2). Integration of all species delimitation methods yielded 15 putative species in the siliculosi group. Two of them corresponded to E. siliculosus and E. crouaniorum, the rest were named using numbers 1 to 13 (Figure 1a).

Ectocarpus putative species consolidation using ITS1 sequences

As reported by [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF], the alignment of all the ITS1 sequences from the siliculosi group was not possible because of their high degree of divergence and the presence of numerous indels. Therefore, alignments were made for four sub-groups that were established based on sequence similarity: (1) Ectocarpus 1, 2, 4 and E. siliculosus (alignment length 885bp); (2) Ectocarpus 5, 6 and 7 (alignment length 447bp); (3) Ectocarpus 8, 9, 10 and 11 (alignment length 456bp); (4) Ectocarpus 12, 13 and E. crouaniorum (alignment length 686 bp). The ITS1 sequences of Ectocarpus 3 did not align well with any other putative species and were not included in these analyses. The topology of the unrooted ITS1 trees built for the alignment sub-groups were congruent with the putative species defined using the COI-5P sequences (Figure 1b). The ABGD analyses conducted within these four sub-groups of alignments were also remarkably congruent with the putative species defined using the COI-5P sequences (Figure 1b, Figure S3). The only discrepancy was the merging of Ectocarpus 12 and Ectocarpus 13 in the ABGD results (Figure 1b). Since both Ectocarpus 12 and Ectocarpus 13 were non-ambiguously defined as two separated putative species with the COI-5P and were retrieved as monophyletic groups in both tree reconstructions for ITS1, we decided to retain 15 consolidated species within the siliculosi group (Figure 1).

Cases of incongruences between markers

Even though results were largely concordant between markers and methods in delineating 15 consolidated species within the siliculosi group, incongruences between the nuclear and the mitochondrial markers were observed in several individuals collected in Chile (Table 2). In particular, a total of 20 individuals collected in Pan de Azúcar (site 26) and 9 individuals collected in Quintay (site 28), were identified as E. crouaniorum using the mitochondrial marker COI-5P but as Ectocarpus 12 using the nuclear marker ITS1. Moreover, one individual sampled in Quintay (site 28), which was identified as Ectocarpus 10 using the COI-5P mitochondrial marker, was identified as Ectocarpus 11 based on the nuclear ITS1 marker and one individual collected in Concepción (site 29), which was identified as Ectocarpus 9 using the COI-5P was also identified as Ectocarpus 11 based on the nuclear marker.

Geographic distribution of the 15 Ectocarpus species

We chose to display the delimitation of the geographic distribution of the 15 species using the results of the mitochondrial COI-5P marker alone. However, it should be noted that a few discrepancies existed between the mitochondrial and the nuclear markers used in this study (see previous paragraph). Distribution patterns varied greatly among the species (Figure 2). Concerning the 10 most commonly sampled species, one seemed to be restricted to a single biogeographic region (Ectocarpus 7, n=32, found only in the Peruvian Province); four species showed a distribution limited to one ocean (Ectocarpus 1, n=30, and Ectocarpus 6, n=48, both found in North and South Pacific; Ectocarpus 10, n=39, distributed only in the South Pacific and Ectocarpus 3, n=29, observed only in the North Atlantic); and five species were encountered in more than one ocean. Among these five species, Ectocarpus 12 (n=16) and Ectocarpus 13 (n=34) were distributed in both the South Pacific and the North Atlantic. In the case of Ectocarpus 8 (n=14), the species was present in both oceans but only one sample was found in the Atlantic. E. siliculosus and E. crouaniorum were the most common species in our data set. E. siliculosus (n=206) was distributed in the Mediterranean Sea, the Pacific and the North Atlantic Oceans, and E. crouaniorum (n=258) was distributed in the South Pacific and the North Atlantic Oceans.

Haplotype networks were created for the 8 species for which >15 sequences were available for both markers (Figure 3). A star-like network pattern, which is usually associated with a recent population expansion, was obtained for six of the eight species studied. For example, E. siliculosus and E. crouaniorum had a single, frequent and widespread haplotype together with several less frequent haplotypes generally restricted to oceanic regions (Northern Atlantic, Lusitanian region, Mediterranean Sea or Peruvian province, Figure 3); this topology was consistent for both COI-5P and ITS1. In contrast, a more reticulate and complex haplotype network was observed for both markers with Ectocarpus 6.

Genetic diversity of E. siliculosus, E. crouaniorum and Ectocarpus 6 Genetic diversity estimates (nH, H, π and S) were generally much lower for the mitochondrial marker than for the nuclear marker (Table S3, S4, S5 for E. siliculosus, E. crouaniorum and Ectocarpus 6, respectively). Regardless of the gene considered, no clear pattern of genetic diversity distribution could be detected within each of the three species (Table S3, S4, S5).

COI-5P data for E. siliculosus (Table S3) indicated that the highest number of haplotypes was on the NW Iberian Peninsula (nH=6, even though close values, i.e. nH=5, were also found for the United Kingdom, France and Chile) whereas the highest values of genetic diversity and nucleotide diversity were in Chile (H= 0.748 +/-0.028 and π =0.237 +/-0.164). ITS1 data gave similar results with the highest value of genetic diversity for the NW Iberian Peninsula (H= 0.862 +/-0.045), the highest nucleotide diversity for Chile (π =0.653 +/-0.364) and the highest number of polymorphic sites for France (S= 19) (Table S3). On the other hand, COI-5P data for E. crouaniorum (Table S4) indicated that the highest number of haplotypes as well as the highest values of nucleotide diversity and polymorphic sites were in France (nH=10, π =0.170 +/-0.130, S= 11, Table S4) while the highest values of genetic diversity were in France and in the NW Iberian Peninsula (H= 0.624 +/-0.092 and 0.679 +/-0.080 respectively). ITS1 data for this same species indicated, however, that the highest number of haplotypes was in the United Kingdom (nH=23) and Chile (nH=23) while all values of genetic diversity were close to one, whatever the region (Table S4). Within Ectocarpus 6, because of the distribution pattern of this species (Figure 3), estimations of genetic diversity were carried out only along the Chilean coast (Table S5). The population of Las Cruces (LAC, Table 1), the largest population of Ectocarpus 6 (n=28), showed the highest number of haplotypes, genetic diversity, and polymorphic sites for both markers: nH= 6, H= 0.791+/-0.048 and S= 8 for COI-5P, and nH= 19, H= 0.934 +/-0.0343 and S= 38 for ITS1 (Table S5).

AMOVA analysis of the cosmopolitan species E. siliculosus and E. crouaniorum

Results of the nested Analysis of Molecular Variance (AMOVA) for both markers are given in Tables 3 and4 for E. siliculosus and E. crouaniorum, respectively. These analyses suggested that the total genetic variance was mainly explained by variance within sites: 55, 81% and 65,18% in E. siliculosus, 59,07% and 77,39% in E. crouaniorum, for the COI-5P and the ITS1, respectively. The variances among regions (< 13% in E. siliculosus and < 11% in E. crouaniorum) and among sites within regions (< 32% in E. siliculosus and < 31% in E. crouaniorum), although significant, were lower than the variance within sites (Tables 3 and4).

Tide level distribution of E. siliculosus, E. crouaniorum and Ectocarpus 6

Ectocarpus species occurred from the upper subtidal up to intertidal pools (Figure 4).

Along the North Atlantic coast, different tide-level distributions were observed for the two most abundant Ectocarpus species (see Figure 4). E. crouaniorum occurred from the high intertidal to the high subtidal but was most abundant within higher intertidal pools (Figure 4). E. siliculosus, which also occurred from the high intertidal to the high subtidal, was most abundant in the lower tidal areas (Figure 4). The distribution of E. crouaniorum followed the same pattern in Chile as on the North Atlantic coast. For E. siliculosus, no clear pattern of distribution could be inferred in Chile since most samples were collected as drifting thalli that had been washed ashore. Ectocarpus 6 was found from medium intertidal to subtidal levels and was more abundant in the low intertidal (Figure 4).

Discussion

In this study, we have characterized the species diversity, geographic distribution, and phylogeographic patterns within the group siliculosi of the genus Ectocarpus. The results presented are based on the most extensive sampling of this group available to date. Using a mitochondrial and a nuclear marker, two complementary species delineation techniques and two tree reconstruction methods, we propose the recognition of 15 putative species within the siliculosi group. E. siliculosus and E. crouaniorum, the only two named species within the siliculosi group (Peters et al. 2010a, 2010b, 2015, Couceiro et al. 2015), were recovered as different species belonging to the most highly divergent clades in our study. We confirmed also that the genome-sequenced species (that we referred as Ectocarpus 7) is different from E. siliculosus, as recently suggested by [START_REF] Peters | Barcoding of cryptic stages of marine brown algae isolated from incubated substratum reveals high diversity in Acinetosporaceae (Ectocarpales, Phaeophyceae)[END_REF]. Moreover, we found individuals showing incongruences between the nuclear and mitochondrial markers suggesting introgression, hybridization or incomplete lineage sorting between some of the newly delineated closely related species. Finally, our extensive sampling along the NEA and SEP coasts revealed that the 15 Ectocarpus species showed different patterns of distribution varying from rare to common cosmopolitan species. Haplotype network topologies for the commonest species showed different patterns of genetic structure suggesting different evolutionary histories.

High species diversity within the siliculosi group

Concordance across results obtained with different methods (monophyly in tree reconstruction, ABGD and GMYC) and the use of unlinked molecular markers (COI-5P and ITS1) are now widely acknowledged methods of supporting the delimitation of previously undescribed species (Carstens et al. 2013, Modica et al. 2014). Indeed, one could expect that unlinked selectively neutral genes will attain concordant genealogical histories when taxa have undergone species-level divergence (i.e., no gene flow for a sufficient amount of time), while reticulate genealogical patterns across those unlinked loci will be observed when genetic exchange exists between taxa [START_REF] Sites | Operational criteria for delimiting species[END_REF]. Our results showed an 80% concordance between methods, a result congruent with studies undertaken in hyper-diverse taxa such as insects [START_REF] Kekkonen | DNA barcode-based delineation of putative species: efficient start for taxonomic workflows[END_REF]. Previous studies have shown that GMYC can lead to an overestimation of group partitioning while ABGD is considered as a more conservative method to delimit species (Puillandre et al. 2012a, 2012b[START_REF] Kekkonen | DNA barcode-based delineation of putative species: efficient start for taxonomic workflows[END_REF], a result concordant with what was observed in the siliculosi group. Despite few discordances detected between our ABGD and GMYC analyses, all putative genetic groups formed highly divergent singletons or monophyletic groups for both markers. For the COI-5P marker, the barcode gap ranged from 0.011-0.037 K2P pairwise genetic distance, which 20 included the cut-off value (0.018) proposed empirically (by eye) by [START_REF] Peters | Barcoding of cryptic stages of marine brown algae isolated from incubated substratum reveals high diversity in Acinetosporaceae (Ectocarpales, Phaeophyceae)[END_REF] for Ectocarpus and other genera of Ectocarpales.

The phylogenetic relationships among the 15 species revealed the occurrence of a monophyletic group composed of E. crouaniorum, Ectocarpus 12, Ectocarpus 13 and a paraphyletic assemblage composed of the remaining 12 other species within the siliculosi group. This branching pattern was retrieved in all previous phylogenetic studies despite a disagreement between the tree topologies depending on the marker used (ITS1 and rubisco spacer: [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF] Incomplete and/or uneven taxon sampling could produce different tree topologies.

Previous studies have reported that the inclusion of additional taxa in a phylogenetic analysis can increase (on average) the accuracy of the inferred topology [START_REF] Lecointre | Species sampling has a major impact on phylogenetic inference[END_REF][START_REF] Hillis | Is sparse taxon sampling a problem for phylogenetic inference?[END_REF][START_REF] Hedtke | Resolution of phylogenetic conflict in large data sets by increased taxon sampling[END_REF].

A problem generated by the various attempts at resolving the Ectocarpus phylogeny is the use of various species codes [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF], Peters et al. 2010a, 2010b).

In our work some clades previously described in the literature were retained as different species (clades 1a, 1c, 2a, 2b, 2c and 3;[START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF], Peters et al. 2010a, 2010b), while other clades were split into different species (1b and 4) [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF], Peters et al. 2010a, 2010b) (see Table S6 for the correspondence between previously distinguished "lineages" and the species code proposed in this study).

Our study detected high levels of cryptic species diversity in the siliculosi group, as suggested in previous studies [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF], Peters et al. 2010a, 2010b). However, it is highly probable that more species exist within this species complex. Indeed, within our data set, two species comprised more than 62% of the samples sequenced (E. siliculosus and E. crouaniorum) while 3 species (Ectocarpus 2, Ectocarpus 4 and Ectocarpus 5) were rare and were represented by less than 5 individuals. It will be necessary to carry out additional population sampling including a better representation of different biogeographic regions in order to better estimate species diversity and distribution of Ectocarpus in a worldwide context. The temperate waters of the southern Australia and the NW Pacific require particular scrutiny since the few sequenced samples from this region (N=12, 1.5% of the samples sequenced) included two species not encountered in other regions (Ectocarpus 2 and Ectocarpus 5).

The high number of cryptic species present in sympatry within the same locality, especially in Chile, raises the question of what evolutionary mechanisms could reduce interspecific competition and promote such patterns. Peters et al. (2010a), Couceiro et al. (2015) and [START_REF] Geoffroy | Molecular evidence for the coexistence of two sibling species in Pylaiella littoralis (Ectocarpales, Phaeophyceae) along the Brittany coast[END_REF] showed that tide level, substratum and season are important factors that have to be taken into account when studying filamentous Ectocarpales. They reported that different species could occupy different spatiotemporal ecological niches related to different tide levels and/or host specificity. For example, Peters et al. (2010a) and [START_REF] Couceiro | Evolution and maintenance of haploid-diploid life cycles in natural populations: the case of the marine brown alga Ectocarpus[END_REF] showed that E. crouaniorum was located higher on the shore than E. siliculosus in NW France. In our study, this difference in tide level distribution between E. crouaniorum and E. siliculosus was corroborated for additional sites in the North Atlantic. Moreover, among the Chilean coast, Ectocarpus 6 seemed more restricted to mid-intertidal pools. These first results provide a good opportunity to study the importance of ecological differentiation between the cryptic species of Ectocarpus.

Incongruence between markers

The huge variability in the level of genetic divergence between species revealed in this study will allow the correlation between reproductive incompatibility and degree of species divergence to be investigated in this genus. Cross-compatibility experiments have been carried out between laboratory strains of E. siliculosus and E. crouaniorum (Peters et al. 2010a), between E. siliculosus and Ectocarpus 7, between Ectocarpus 7 and Ectocarpus 1 [START_REF] Peters | Proposal of Ectocarpus siliculosus (Ectocarpales, Phaeophyceae) as a model organism for brown algal genetics and genomics[END_REF], and between E. siliculosus and Ectocarpus 1 [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF]. Hybrid sporophytes from the four crosses were viable but incapable of meiosis. [START_REF] Müller | Sexual reproduction of Ectocarpus siliculosus (Ectocarpales, Phaebphyceae) in Japan[END_REF], in contrast, crossed an Ectocarpus from Japan, which is closely related to the genome-sequenced strain based on its ITS sequence but for which COI-5P sequences are so far unavailable, with E. siliculosus, and obtained meiosis-competent hybrid sporophytes. This cross definitively needs confirmation.

The situation in the field is less well studied. Peters et al. (2010aPeters et al. ( , 2010b) ) revealed the presence of field hybrids between E. siliculosus and E. crouaniorum in Chile and France but nothing is known about the proportion of hybrids in natural populations.

Inspection of tree topologies obtained for several loci from different compartments have been successfully applied to identify potential cases of introgression and ancient hybridization events in natural populations [START_REF] Peters | Nuclear loci and coalescent methods support ancient hybridization as cause of mitochondrial paraphyly between gadwall and falcated duck (Anas spp.)[END_REF]). In our data set, a low percentage of incongruence (6%) was observed between results obtained with the mitochondrial and the nuclear marker. The species involved were phylogenetically more closely related than the crosses mentioned in the last paragraph (Figure 1a,1b).

Incongruent individuals were found exclusively among Chilean samples collected at sites where the respective species were in contact. Taken together, our results suggest the existence of different levels of reproductive barriers within the E. siliculosi complex, leading to mtDNA introgression only between some species pairs. Incomplete reproductive isolation may have an important bearing on the evolutionary trajectories of species by decreasing divergence between species but also by allowing new favorable mutations and allelic combinations to transgress species boundaries [START_REF] Allendorf | The problems with hybrids: setting conservation guidelines[END_REF][START_REF] Mallet | Hybridization as an invasion of the genome[END_REF]. However, both incomplete lineage sorting and hybridization lead to similar gene tree incongruence signatures and distinguishing between those two processes has proven difficult [START_REF] Knowles | The burgeoning field of statistical phylogeography[END_REF]). Hybridization may be high between closely related species, unfortunately, incomplete lineage sorting is also likely to be at least partly responsible of the gene tree incongruences in species complexes of recent origin. New statistical frameworks, which allow testing for hybridization despite incomplete lineage sorting, have been developed recently (see Yu et al. 2011 and references therein). More extensive sampling, adapted genetic tools and analyses are needed to estimate the extent and importance of hybridization between the species of the siliculosi group in the field.

Species distribution

During our study we extensively sampled two coasts where strong biogeographic boundaries are recognized. Along the Chilean coast, two biogeographic boundaries have been described [START_REF] Camus | Biogeografía marina de Chile continental[END_REF]). The first is located at 30-33°S and separates the Peruvian Province from the Intermediate Area; the second is located at 42°S and separates the Intermediate area from the Magellanic Province. Along the European coast, the Celtic-Sea/Brittany area has been described as a biogeographical transition zone between the Northern European Sea and the Lusitanian Province while the front Almería-Oran separates the Mediterranean coasts from the Atlantic ones [START_REF] Spalding | Marine ecoregions of the world: a bioregionalization of coastal and shelf areas[END_REF]. Several phylogeographic studies have reported a concordance between genetic discontinuities and biogeographic boundaries, attributing this pattern to the existence of historical barriers caused by oceanographic or climatic features (for Chile see the reviews [START_REF] Haye | Phylogeographic structure in benthic marine invertebrates of the southeast Pacific coast of Chile with differing dispersal potential[END_REF][START_REF] Guillemin | Phylogeography of seaweeds in the South East Pacific: complex evolutionary processes along a latitudinal gradient[END_REF]; for Europe see see the reviews [START_REF] Maggs | Evaluating signatures of glacial refugia for North Alantic benthic marine taxa[END_REF][START_REF] Neiva | Climate oscillations, range shifts and phylogeographic patterns of North Atlantic Fucaceae[END_REF]. Interestingly, the species for which the genome has been sequenced, Ectocarpus 7 (n=32), is apparently restricted to the Peruvian Province; Peters et al. (2010b) previously found similar results using a smaller sampling scheme. The Peruvian Province is characterized by continuous upwelling of cool water (16-20°C at the sea surface) and is also affected by recurrent El Niño events, causing several weeks of higher sea surface temperatures (more than 10°C of amplitude) as a result of the southward incursion of warm waters [START_REF] Peters | Temperature tolerance and latitudinal range of brown algae from temperate Pacific South America[END_REF]. The distribution range of Ectocarpus 7 may reflect an adaptation of individuals to this specific oceanographic environment. Apart for Ectocarpus 7, our results did not support the existence of extensive biogeographic or phylogeographic breaks for other Ectocarpus species. This lack of phylogeographic structure has been reported for species that have high dispersal capacities and/or for invasive species [START_REF] Cárdenas | A phylogeographical analysis across three biogeographical provinces of the south-eastern Pacific: the case of the marine gastropod Concholepas concholepas[END_REF][START_REF] Haye | Phylogeographic structure in benthic marine invertebrates of the southeast Pacific coast of Chile with differing dispersal potential[END_REF][START_REF] Guillemin | Tracing the trans-Pacific evolutionary history of a domesticated seaweed (Gracilaria chilensis) with archaeological and genetic data[END_REF]). Short dispersal distances of spores, gametes or zygotes coupled with rare events of long distance colonization seem to be the rule for the Phaeophyceae [START_REF] Reed | The effects of variable settlement and early competition on patterns of kelp recruitment[END_REF][START_REF] Raimondi | Effects of selffertilization in the giant kelp, Macrocystis pyrifera[END_REF][START_REF] Neiva | Fine-scale genetic breaks driven by historical range dynamics and ongoing density-barrier effects in the estuarine seaweed Fucus ceranoides L[END_REF][START_REF] Robuchon | Contrasting genetic diversity patterns in two sister kelp species co-distributed along the coast of Brittany[END_REF]. However, Ectocarpus is described as an important contributor to biofouling and long-distance dispersal might be associated with human transport [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF]. Indeed, we detected two truly cosmopolitan species (E.

siliculosus and E. crouaniorum) showing a star-like haplotype network and shared haplotypes between continents, a pattern characteristic of a recent expansion that may be facilitated by human activities. Ectocarpus 1 and Ectocarpus 6 occur in both the northern and the southern Pacific Ocean. Despite the large distances separating the sampled populations of these two species, no genetic structure was detected in their haplotype networks, a pattern suggestive of recent dispersal events between the North and South Pacific coasts. Similarly, no phylogeographic structure was observed for three South Pacific species (Ectocarpus 9, Ectocarpus 10 and Ectocarpus 11) nor for one North Atlantic species (Ectocarpus 3). It has been suggested that recent colonization events can eliminate genetic structure linked to historical barriers [START_REF] Smith | The role of historical and contemporary processes on phylogeographic structure and genetic diversity in the Northern Cardinal, Cardinalis cardinalis[END_REF][START_REF] Dibattista | Phylogeography of two closely related Indo-Pacific butterflyfishes reveals divergent evolutionary histories and discordant results from mtDNA and microsatellites[END_REF], thus the introductions of Ectocarpus species through shipping activities could explain the lack of phylogeographic structure in our study. [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF] have indeed reported that strains belonging to E. siliculosus sensu stricto (referred as lineage 1a) sampled from different continents show a maximum of 5 substitutions for the ITS marker; these authors suggest that recent dispersal events could have shaped the genetic diversity in this species. Both natural dispersal after the Pleistocene and transport via shipping have been proposed for this species [START_REF] Stache-Crain | Molecular systematics of Ectocarpus and Kuckuckia (Ectocarpales, Phaeophyceae) inferred from phylogenetic analyses of nuclear and plastid-encoded DNA sequences[END_REF]). In our study, the distribution of Ectocarpus 8 seems related to dispersal associated with human transport. This species was present in the South Pacific and a single sample was found in the North Atlantic. This sample corresponds to an individual collected in the Kingsbridge Estuary (Devon, England). This could suggest a recent arrival of this species in the English Channel in ballast water or attached to a ship hull. Even if the dispersal capacity of Ectocarpus might be favored by shipping, when the number of samples was sufficient to perform within-species genetic differentiation analyses, we always found a slight but significant hierarchical pattern of genetic differentiation (i.e. see Table 3 and4; results of the nested AMOVA).

Consequently, the pattern of genetic differentiation may be more complex than what we found in this study and sampling effort needs to be improved for all species in order to get a comprehensive idea of species distribution, species phylogeography and population connectivity.

Conclusion

Using DNA sequence data and species delimitation methods we have observed the presence of at least fifteen species within the Ectocarpus siliculosi group. Species showed different patterns of distribution and suggested different evolutionary histories.

Further scrutiny of individuals cultivated in controlled laboratory conditions may reveal consistent morphological differences between species. However, future research on speciation in these filamentous brown algae will have to take into account that in the field, it is impossible to distinguish between species within the E. siliculosi group. This contrasts clearly with the significant morphological differences observed between recently diverging species in other Phaeophyceae, such as Fucus [START_REF] Coyer | Evolution and diversification within the intertidal brown macroalgae Fucus spiralis/F. vesiculosus species complex in the North Atlantic[END_REF][START_REF] Cánovas | Driving south: a multi-gene phylogeny of the brown algal family Fucaceae reveals relationships and recent drivers of a marine radiation[END_REF]). In any case, the complex pattern of phylogenetic relationships among the 15 species revealed in this study, opens a very interesting field of research deciphering the process of evolution and diversification in this group using the tools available from the model organism for genomics and genetics of the brown macroalgae. Table 1. Sites and samples sequenced in the present study. The number of site, the continent, country, name of site, code, year of sampling and 993 the number of sequences for both markers (COI-5P and ITS1) are indicated. 994 according to the mitochondrial marker COI-5P. Sites are numbered as in Table 1.

N°

  ; ITS1, ITS2, Rubisco spacer region, cox3 and rps14-atp8: Peters et al. 2010a; ITS1, ITS2, Rubisco spacer and cox3: Peters et al. 2010b, COI-5P: Peters et al. 2015; COI-5P and ITS1: this study).

  a doctoral grant to A. E. Montecinos (Becas-Chile, CONICYT, advanced human resources program), additional support from the projects IDEALG (France: ANR-10-BTBR-04), "Bibliothèque du vivant" (France: INRA-MNHN-INEE-CNRS) and Bi-Cycle (France: ANR10-BLAN-1727), the EU FP7 "capacities" specific program ASSEMBLE (grant no. 227799), the EU Interreg program France (Channel)-England (project Marinexus), the TOTAL Foundation (Project "Brown algal biodiversity and ecology in the Eastern Mediterranean Sea"), and a Ray-Lankester Fellowship (UK: Marine Biological Association, Plymouth) to A. F. Peters. The work was carried out within the context of the international research network "Diversity, Evolution and Biotechnology of Marine Algae" (GDRI N° 0803). The authors have declared that no competing interests exist. Tables 992

  Figure 1. Proposed species in the Ectocarpus siliculosi group using COI-5P (a) and their

Figure 2 .

 2 Figure 2. Distribution of the 15 species of the Ectocarpus subgroup siliculosi as defined

Figure 3 .

 3 Figure 3. Haplotype networks of for COI-5P and ITS1 for eight Ectocarpus spp..

Figure 4 .

 4 Figure 4. Distribution on the shore of the commonest Ectocarpus species in the North

Figure S1 .Figure S2 .

 S1S2 Figure S1. Distribution of pairwise distances for the marker COI-5P and automatic 1

-

  Figure S3. Distribution of pairwise distances between haplotype pairs for the marker 1

Table 2 .

 2 Association between mitochondria (COI-5P) and nrDNA (ITS1) sequences in the Ectocarpus specimens in which both markers were sequenced. Individuals showing incongruence among markers are indicated in bold. Esil = E. siliculosus, Ecro = E. crouaniorum, Ec 1 -12 = Ec 2 Ec 3 Ec 4 Ec 5 Ec 6 Ec 7 Ec 8 Ec 9 Ec 10 Ec 11 Ec 12 Ec 13

	of site	Continent	Country	Site	CODE	YEAR	COI-5P ITS1
	1	Europe	UK	Wick	WIC	2008	10	10
	2	Europe	UK	Rattray Head	RAT	2008	1	0
	3	Europe	UK	Dunstaffnage	DUN	2008	7	8
	4	Europe	UK	Berwick	BER	2008	12	12
	5	Europe	UK	Mull of Galloway	MUL	2008	14	14
	6	Europe	UK	Pett level	PET	2008	11	13
	7	Europe	UK	Gosport Marina	GOS	2008	1	3
	8	Europe	UK	Plymouth	PLY	2010-2011	38	37
	9	Europe	UK	Restronguet	RES	2010	13	12
	10	Europe	France	Cherbourg	CHE	2006	2	0
	11	Europe	France	Roscoff	ROS	2010-2012	48	31
	12	Europe	France	Saint Malo	STM	2010	23	0
	13	Europe	France	Traezh Hir	THZ	2010	28	24
	14	Europe	France	Concarneau	CON	2010	1	0
	15	Europe	France	Quiberon	QUI	2010-2012	30	27
	16	Europe	Spain	Ribadeo	RIB	2013	27	25
	17	Europe	Spain	Coruña	COR	2013	26	19
	18	Europe	Spain	Ría de Arousa	RIA	2013	8	8
	19	Europe	Portugal	Viana	VIA	2013	11	8
	20	Europe	Italy	Naples	NAP	2012	42	27

Table 3 .

 3 Analysis of molecular variance (AMOVA) of E. siliculosus for each molecular marker (COI-5P and ITS1). Regions, not including the 999 Mediterranean for which only a single site was sampled, as in Supplementary Table3. d.f.: degree of freedom. SS: sum of squares. 1000

	Source of variation	d.f.	SS	Variance components % Variation p-value
	COI-5P
	Among regions	3	18.982	0.07307	12.54	<0.0001
	Among sites within regions	10	22.304	0.18446	31.65	<0.0001
	Within site	154	50.095	0.32529	55.81	0.0351
		TOTAL 167	91.381	0.58283
	ITS1
	Among regions	3	51.486	0.15194	7.14	<0.0001
	Among sites within regions	10	74.673	0.58877	27.68	<0.0001
	Within site	154 213.502	1.38638	65.18	0.1476
		TOTAL 167 339.661	2.12709

Table 4 .

 4 Analysis of molecular variance (AMOVA) of E. crouaniorum for each molecular marker (COI-5P and ITS1). Regions as in 1001 Supplementary Table4. d.f.: degree of freedom. SS: sum of squares. 1002

	Source of variation	d.f.	SS	Variance components % Variation p-value
	COI-5P
	Among regions	3	11.404	0.05270	10.74	<0.0001
	Among sites within regions	12	19.118	0.14818	30.19	<0.0001
	Within site	134	38.851	0.28993	59.07	0.0088
		TOTAL 149	69.373	0.49082
	ITS1
	Among regions	3	37.469	0.08576	2.67	<0.0001
	Among sites within regions	12	97.254	0.63927	19.93	<0.0001
	Within site	134 332.610	2.48217	77.39	0.37634
		TOTAL 149	69.373	0.49082

Table S1 .

 S1 COI-5P sequences used in the present study. Details of the date, geographic origin, Genbank accession number and reference 1

		WIC08-04 EcQAB10-02 EcPLY10-23 BLZ11-51 L213 EcNAP12-120 EcNAP12-s3-50 EcPAN-174 EcBIN-294 EcBIN-09 EcCIS-073 EcQUI-509 EcLAC-403 EcLAC-415 EcCOL-559 EcCNA-659 EcEST-795	United Kingdom United Kingdom 2010 United Kingdom 2011 France France Italy 2012 Italy Spain Chile 2013 Chile 2013 Chile 2013 Chile Chile 2013 Chile 2013 Chile Chile 2013 Chile	Wick Plymouth Plymouth Roscoff Saint Malo Naples Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Caleta Cisne, Caldera Quintay Las Cruces Las Cruces Cocholgüe, Tomé Curiñanco, Valdivia Estaquilla	KT983134 KT982765 KT983095 KT982831 KT983396 KT982812 KU134153 KT982873 KT982889 KU134021 KT987931 KT982988 KT982973 KT987952 KT987969 KT983023 KT983038	This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper
	2	of each COI-5P sequences used in the present study. N° Name of sample Date Country 1 SAM08-3 2008 United Kingdom 2 GOS08-02 2008 United Kingdom 3 Ec318 2006 France 4 BLZ11-27 2011 France 5 BLZ11-78 2011 France 6 BLZ11-47 2011 France 7 EcVAZ11-10 2011 France 8 EcNAP12-64 2012 Italy 9 GR11-33 2011 Greece 10 GR11-38B 2011 Greece 11 GR11-12A 2011 Greece 12 Ec294 2006 Peru 24 WIC08-05 United Kingdom 25 WIC08-07 United Kingdom 26 WIC08-08 United Kingdom 27 WIC08-17 United Kingdom 28 WIC08-18 United Kingdom 29 WIC08-19 United Kingdom 30 RAT08-6 United Kingdom 31 BUT08-05 United Kingdom 32 BUT08-11 United Kingdom 33 BUT08-13 United Kingdom 34 BUT08-24 United Kingdom 35 BUT08-25 United Kingdom 36 BUT08-26 United Kingdom 37 BUT08-27 United Kingdom 38 BUT08-30 United Kingdom 76 EcQAB10-03 United Kingdom 77 EcQAB10-04 United Kingdom 78 EcQAB10-05 United Kingdom 79 EcQAB10-06 United Kingdom 80 EcQAB10-07 United Kingdom 81 EcQAB10-09 United Kingdom 82 EcQAB10-11 United Kingdom 83 EcQAB10-12 United Kingdom 84 EcQAB10-13 United Kingdom 85 EcQAB10-14 United Kingdom 86 EcQAB10-15 United Kingdom 87 EcQAB10-16 United Kingdom 88 EcQAB10-17 United Kingdom 89 EcQAB10-18 United Kingdom 90 EcQAB10-19 United Kingdom 102 EcPLY10-25 2010 United Kingdom 103 EcPLY10-26 2010 United Kingdom 104 EcPLY10-27 2010 United Kingdom 105 EcPLY10-28 2010 United Kingdom 106 EcPLY10-29 2010 United Kingdom 107 EcPLY10-30 2010 United Kingdom 108 EcPLY10-31 2010 United Kingdom 109 EcPLY10-32 2010 United Kingdom 110 EcPLY10-34 2010 United Kingdom 111 Ec242 2011 United Kingdom 112 EcREP10-38 2010 United Kingdom 113 EcREP10-39 2010 United Kingdom 114 EcREP10-42 2010 United Kingdom 115 EcREP10-43 2010 United Kingdom 116 EcREP10-44 2010 United Kingdom 154 EcPH11-s-2A-09 2011 France 155 EcPH11-s-2A-16 2011 France 156 EcPH11-s2A-01 2011 France 157 EcPH11-s2A-14 2011 France 158 EcPH11-21 2011 France 159 EcPH11-16 2011 France 160 EcPH11-09 2011 France 161 BLZ11-44 2011 France 162 EcPH11-s5-38 2011 France 163 EcPH11-s5-07 2011 France 164 EcPH11-38 2011 France 165 EcPH11-33 2011 France 166 EcPH11-28 2011 France 167 EcPH11-27 2011 France 168 EcPH11-47 2011 France 180 L215 France 181 L209 France 182 L210 France 183 L195 France 184 L197 France 185 L198 France 186 L205 France 187 L190 France 188 L183 France 189 L170 France 190 L216 France 191 L168 France 192 L196 France 193 EcTH10-04 France 194 EcTH10-05 France 336 EcNAP12-210 Italy 337 EcNAP12-s3-34 Italy 338 EcNAP12-15 Italy 339 EcNAP12-17 Italy 340 EcNAP12-14 Italy 341 EcNAP12-12 Italy 342 EcNAP12-94 Italy 343 EcNAP12-s3-17 Italy 344 EcNAP12-s3-11 Italy 345 EcNAP12-s3-13 Italy 346 EcNAP12-s3-19 Italy 347 EcNAP12-s3-22 Italy 348 EcNAP12-s3-23 Italy 349 EcNAP12-s4-16 Italy 350 EcNAP12-s4-18 Italy 362 EcNAP12-s4-19 2012 Italy 363 EcNAP12-s4-09 2012 Italy 364 EcNAP12-82 2012 Italy 365 GR11-28 2011 Greece 366 GR11-36 2011 Greece 367 GR11-38A 2011 Greece 368 GR11-40 2011 Greece 369 GR11-45 2011 Greece 370 GR11-46 2011 Greece 371 GR11-48 2011 Greece 372 LES08 2009 Greece 373 LES15 2009 Greece 374 LES11 2009 Greece 375 LES4 2009 Greece 376 LES1 2009 Greece 388 EcPAN-175 Spain 389 EcPAN-192 Spain 390 EcPAN-197 Spain 391 EcPAN-161A Chile 392 EcPAN-117 Chile 393 EcPAN-119 Chile 394 EcPAN-177 Chile 395 EcPAN-183 Chile 396 EcPAN-185 Chile 397 EcPAN-233 Chile 398 EcPAN-083 Chile 399 EcPAN-087A Chile 400 EcPAN-091A Chile 401 EcPAN-097A Chile 402 EcPAN-085 Chile 440 EcBIN-295 Chile 441 EcBIN-306 Chile 442 EcBIN-307 Chile 443 EcBIN-308 Chile 444 EcBIN-309 Chile 445 EcBIN-315 Chile 446 EcBIN-320 Chile 447 EcBIN-323 Chile 448 EcBIN-330 Chile 449 EcBIN-331 Chile 450 EcBIN-332 Chile 451 EcBIN-333 Chile 452 EcBIN-337 Chile 453 EcBIN-338 Chile 454 EcBIN-339 Chile 466 EcBIN-10 2013 Chile 467 EcBIN-220 2013 Chile 468 EcBIN-221 2013 Chile 469 EcBIN-224 2013 Chile 470 EcBIN-226 2013 Chile 471 EcBIN-243 2013 Chile 472 EcBIN-246 2013 Chile 473 EcBIN-247 2013 Chile 474 EcBIN-276 2013 Chile 475 EcBIN-346 2013 Chile 476 EcBIN-348 2013 Chile 477 EcBIN-350 2013 Chile 478 EcBIN-351 2013 Chile 479 EcBAS-252A 2013 Chile 480 EcBAS-264 2013 Chile EcPVI-36 2013 Chile EcPVI-035 2013 Chile Ec454 ---Chile EcALG-520 2013 Chile EcALG-521 2013 Chile EcALG-522 2013 Chile EcALG-523 2013 Chile EcALG-525 2013 Chile EcALG-527 2013 Chile EcALG-521A 2013 Chile EcALG-524 2013 Chile EcQUI-482 2013 Chile EcQUI-483 2013 Chile EcQUI-485 2013 Chile EcQUI-486 2013 Chile 518 EcQUI-510 2013 Chile 519 EcQUI-511 2013 Chile 520 EcQUI-515 2013 Chile 521 EcQUI-516 2013 Chile 522 EcQUI-517 2013 Chile 523 EcQUI-518 2013 Chile 524 EcQUI-455 2013 Chile 525 EcQUI-456 2013 Chile 526 EcQUI-457 2013 Chile 527 EcQUI-467 2013 Chile 528 EcQUI-469 2013 Chile 529 EcQUI-476 2013 Chile 530 EcQUI-477 2013 Chile 531 EcQUI-478 2013 Chile 532 EcQUI-480 2013 Chile 544 EcLAC-397 Chile 545 EcLAC-412 Chile 546 EcLAC-398 Chile 547 EcLAC-424 Chile 548 EcLAC-384 Chile 549 EcLAC-354 Chile 550 EcLAC-355 Chile 551 EcLAC-356 Chile 552 EcLAC-357 Chile 553 EcLAC-363 Chile 554 EcLAC-365 Chile 555 EcLAC-366 Chile 556 EcLAC-369 Chile 557 EcLAC-371 Chile 558 EcLAC-373 Chile 570 EcLAC-425 2013 Chile 571 EcLAC-430 2013 Chile 572 EcLAC-433 2013 Chile 573 EcLAC-435 2013 Chile 574 EcLAC-443 2013 Chile 575 EcLAC-444 2013 Chile 576 EcLAC-445 2013 Chile 577 EcLAC-402A 2013 Chile 578 EcLAC-379 2013 Chile 579 EcDIC-605 2013 Chile 580 EcDIC-577 2013 Chile 581 EcDIC-578 2013 Chile 582 EcDIC-579 2013 Chile 583 EcDIC-581 2013 Chile 584 EcDIC-583 2013 Chile 622 EcCOL-567 2013 Chile 623 EcCOL-568A 2013 Chile 624 EcPIL-611 2013 Chile 625 EcPIL-616 2013 Chile 626 EcPIL-618 2013 Chile 627 EcPIL-624 2013 Chile 628 EcPIL-625 2013 Chile 629 EcPIL-628 2013 Chile 630 EcPIL-650 2013 Chile 631 EcPIL-621 2013 Chile 632 EcPIL-631 2013 Chile 633 EcPIL-632 2013 Chile 634 EcPIL-633 2013 Chile 635 EcPIL-634 2013 Chile 636 EcPIL-635 2013 Chile 648 EcCNA-642 Chile 649 EcSCA-726 Chile 650 EcSCA-721 Chile 651 EcSCA-722 Chile 652 EcSCA-723 Chile 653 EcPLG-708 Chile 654 EcPLG-712 Chile 655 EcPLG-716 Chile 656 EcPLG-717 Chile 657 EcPLG-718 Chile 658 EcPLG-682A Chile 659 EcPLG-686 Chile 660 EcPLG-688 Chile 661 EcPLG-713 Chile 662 EcPLG-714 Chile 674 EcEST-796 2013 Chile 675 EcEST-797 2013 Chile 676 EcEST-798 2013 Chile 677 EcEST-800 2013 Chile 678 EcEST-801 2013 Chile 679 EcEST-802 2013 Chile 680 EcEST-804 2013 Chile 681 EcEST-805 2013 Chile 682 EcEST-806 2013 Chile 683 EcEST-807 2013 Chile 684 EcEST-808 2013 Chile 685 EcEST-809 2013 Chile 686 EcEST-810 2013 Chile 687 EcEST-811 2013 Chile 688 EcEST-813 2013 Chile 97 HAS08-18 United Kingdom 98 HAS08-7 United Kingdom 99 HAS08-8 United Kingdom 100 HAS08-11 United Kingdom 101 HAS08-12 United Kingdom 102 HAS08-13 United Kingdom 103 HAS08-14 United Kingdom 104 HAS08-16 United Kingdom 105 HAS08-17 United Kingdom 106 HAS08-19 United Kingdom 107 HAS08-20 United Kingdom 108 LH14 United Kingdom 109 LH7 United Kingdom 110 LH13 United Kingdom 111 EcQAB10-1 United Kingdom 122 EcQAB10-12 2010 United Kingdom 123 EcQAB10-13 2010 United Kingdom 124 EcQAB10-14 2010 United Kingdom 125 EcQAB10-15 2010 United Kingdom 126 EcQAB10-16 2010 United Kingdom 127 EcQAB10-17 2010 United Kingdom 128 EcQAB10-18 2010 United Kingdom 129 EcQAB10-19 2010 United Kingdom 130 EcQAB10-20 2010 United Kingdom 131 EcQAB10-21 2010 United Kingdom 132 EcQAB10-22 2010 United Kingdom 133 EcQAB10-23 2010 United Kingdom 134 EcQAB10-24 2010 United Kingdom 135 EcPLY10-31 2010 United Kingdom 136 EcPLY10-19 2010 United Kingdom 147 Ec242 United Kingdom 148 EcREP10-38 United Kingdom 149 EcREP10-39 United Kingdom 150 EcREP10-42 United Kingdom 151 EcREP10-43 United Kingdom 152 EcREP10-44 United Kingdom 153 EcREP10-45 United Kingdom 154 EcREP10-48 United Kingdom 155 EcREP10-49 United Kingdom 156 EcREP10-50 United Kingdom 157 EcREP10-51 United Kingdom 158 EcREP10-52 United Kingdom 159 EcREP10-54 United Kingdom 160 Ec334 France 161 EcTH10-10 France 198 EcTH10-12 France 199 EcTH10-13 France 200 EcTH10-14 France 201 EcTH10-15 France 202 EcTH10-16 France 203 EcTH10-17 France 204 EcTH10-18 France 205 EcTH10-53 France 206 EcTH10-26 France 207 EcTH10-31 France 208 EcTH10-32 France 209 EcTH10-33 France 210 EcTH10-35 France 247 EcRIB-195 Spain 248 EcRIB-200 Spain 249 EcRIB-203 Spain 250 EcRIB-204 Spain 251 EcRIB-207 Spain 252 EcRIB-208 Spain 253 EcRIB-231 Spain 254 EcRIB-234 Spain 255 EcRIB-235 Spain 256 EcRIB-236 Spain 257 EcRIB-159 Spain 258 EcRIB-185 Spain 259 EcRIB-187 Spain 260 EcRIB-190 Spain 297 EcVIA-39 Portugal 298 EcPOV-554 Portugal 299 EcVIA-31 Portugal 300 EcVIA-33 Portugal 301 EcVIA-45 Portugal 302 EcNAP12-109 Italy 303 EcNAP12-110 Italy 304 EcNAP12-112 Italy 305 EcNAP12-114 Italy 306 EcNAP12-117 Italy 307 EcNAP12-118 Italy 308 EcNAP12-119 Italy 309 EcNAP12-111 Italy 310 EcNAP12-113 Italy 347 EcPAN-128 Chile 348 EcPAN-138 Chile 349 EcPAN-140 Chile 350 EcPAN-174 Chile 351 EcPAN-175 Chile 352 EcPAN-192 Chile 353 EcPAN-197 Chile 354 EcPAN-104 Chile 355 EcPAN-089 Chile 356 EcPAN-148 Chile 357 EcPAN-161 Chile 358 EcPAN-163 Chile 359 EcPAN-143A Chile 360 EcPAN-149A Chile 372 EcPAN-160 2013 Chile 373 EcPAN-087A 2013 Chile 374 EcPAN-114A 2013 Chile 375 EcPAN-139A 2013 Chile 376 EcPAN-139B 2013 Chile 377 EcPAN-144A 2013 Chile 378 EcPAN-181 2013 Chile 379 EcPAN-182 2013 Chile 380 EcPAN-174A 2013 Chile 381 EcBIN-293 2013 Chile 382 EcBIN-272 2013 Chile 383 EcBIN-273 2013 Chile 384 EcBIN-277 2013 Chile 385 EcBIN-279 2013 Chile 397 EcBIN-309 Chile 398 EcBIN-315 Chile 399 EcBIN-320 Chile 400 EcBIN-323 Chile 401 EcBIN-330 Chile 402 EcBIN-331 Chile 403 EcBIN-332 Chile 404 EcBIN-333 Chile 405 EcBIN-338 Chile 406 EcBIN-339 Chile 407 EcBIN-341 Chile 408 EcBIN-343 Chile 409 EcBIN-010 Chile 410 EcBIN-220 Chile 422 EcBIN-351 2013 Chile 423 EcBIN-224 2013 Chile 424 EcBIN-226 2013 Chile 425 EcBIN-243 2013 Chile 426 EcBIN-247 2013 Chile 427 EcBIN-246 2013 Chile 428 EcBIN-009 2013 Chile 429 EcBAS-255D 2013 Chile 430 EcBAS-298 2013 Chile 431 EcBAS-304 2013 Chile 432 EcBAS-302A 2013 Chile 433 EcBAS-303 2013 Chile 434 EcBAS-305 2013 Chile 435 EcBAS-255A 2013 Chile 447 EcALG-525 Chile 448 EcALG-527 Chile 449 EcALG-524 Chile 450 EcQUI-478 Chile 451 EcQUI-455 Chile 452 EcQUI-467 Chile 453 EcQUI-469 Chile 454 EcQUI-480 Chile 455 EcQUI-456 Chile 456 EcQUI-457 Chile 457 EcQUI-476 Chile 458 EcQUI-477 Chile 459 EcQUI-450 Chile 460 EcQUI-484 Chile 472 EcLAC-369 2013 Chile 473 EcLAC-355 2013 Chile 474 EcLAC-392 2013 Chile 475 EcLAC-366 2013 Chile 476 EcLAC-402A 2013 Chile 477 EcLAC-430 2013 Chile 478 EcLAC-445 2013 Chile 479 EcLAC-354 2013 Chile 480 EcLAC-371 2013 Chile 481 EcLAC-386 2013 Chile 482 EcLAC-444 2013 Chile 483 EcLAC-357 2013 Chile 484 EcLAC-365 2013 Chile 485 EcLAC-407 2013 Chile EcPH10-22 197 211 EcTH10-36 France 261 EcRIB-191 Spain 311 EcNAP12-s4-09 Italy 361 EcPAN-083 Chile 386 EcBIN-283 2013 Chile 411 EcBIN-221 Chile 436 EcBAS-264 2013 Chile 461 EcQUI-490 Chile 486 EcLAC-373 2013 Chile 112 EcQAB10-2 United Kingdom 137 EcPLY10-20 2010 United Kingdom 212 EcTH10-38 France 262 EcRIB-193 Spain 312 EcNAP12-s4-10 Italy 362 EcPAN-085 Chile 387 EcBIN-284 2013 Chile 412 EcBIN-001 Chile 437 EcBAS-305B 2013 Chile 462 EcQUI-499 Chile 487 EcLAC-374 2013 Chile 13 Ec32(genome strain) ---Peru 39 BUT08-31 United Kingdom 91 EcQAB10-20 United Kingdom 117 EcREP10-45 2010 United Kingdom 169 EcPH12-76 2012 France 195 EcTH10-06 France 351 EcNAP12-s4-11 Italy 377 LES2 2009 Greece 403 EcPAN-086 Chile 455 EcBIN-341 Chile 481 EcBAS-298 2013 Chile EcQUI-489 2013 Chile 533 EcQUI-484 2013 Chile 559 EcLAC-374 Chile 585 EcDIC-584 2013 Chile 637 EcPIL-636 2013 Chile 663 EcEST-782 Chile 689 EcEST-815 2013 Chile 113 EcQAB10-3 United Kingdom 138 EcPLY10-21 2010 United Kingdom 213 EcTH10-39 France 263 EcRIB-213 Spain 313 EcNAP12-s4-11 Italy 363 EcPAN-086 Chile 388 EcBIN-285 2013 Chile 413 EcBIN-002 Chile 438 EcBAS252A 2013 Chile 463 EcQUI-500 Chile 488 EcLAC-388 2013 Chile 14 Ec156 2006 Chile 40 BUT08-35 United Kingdom 92 EcQAB10-21 United Kingdom 118 EcREP10-46 2010 United Kingdom 170 L212 2010 France 196 EcTH10-07 France 352 EcNAP12-s3-21 Italy 378 LES3 2009 Greece 404 EcPAN-087 Chile 456 EcBIN-343 Chile 482 EcBAS-303 2013 Chile EcQUI-491 2013 Chile 534 EcQUI-490 2013 Chile 560 EcLAC-386 Chile 586 EcDIC-586 2013 Chile 638 EcPIL-651 2013 Chile 664 EcEST-785 Chile 690 EcEST-816 2013 Chile 114 EcQAB10-4 United Kingdom 139 EcPLY10-23 2010 United Kingdom 214 EcTH10-41 France 264 EcRIB-214 Spain 314 EcNAP12-s4-16 Italy 364 EcPAN-087 Chile 389 EcBIN-288 2013 Chile 414 EcBIN-004 Chile 439 EcCIS-073 2013 Chile 464 EcQUI-501 Chile 489 EcLAC-387 2013 Chile 15 Ec157 2006 Chile 41 BUT08-40 United Kingdom 93 EcQAB10-22 United Kingdom 119 EcREP10-48 2010 United Kingdom 171 L206 2010 France 197 EcTH10-08 France 353 EcNAP12-s3-01 Italy 379 LES7 2009 Greece 405 EcPAN-114A Chile 457 EcBIN-227 Chile 483 EcBAS-304 2013 Chile EcQUI-493 2013 Chile 535 EcQUI-499 2013 Chile 561 EcLAC-387 Chile 587 EcDIC-587 2013 Chile 639 EcPIL-648 2013 Chile 665 EcEST-786 Chile 691 EcEST-769 2013 Chile 115 EcQAB10-5 United Kingdom 140 EcPLY10-26 2010 United Kingdom 215 EcQB10-20 France 265 EcRIB-215 Spain 315 EcNAP12-s4-18 Italy 365 EcPAN-091A Chile 390 EcBIN-291 2013 Chile 415 EcBIN-005 Chile 440 EcCIS-070 2013 Chile 465 EcQUI-504 Chile 490 EcLAC-389 2013 Chile 16 Ec666 2009 USA 42 BUT08-41 United Kingdom 94 EcQAB10-23 United Kingdom 120 EcREP10-49 2010 United Kingdom 172 L211 2010 France 198 EcTH10-09 France 354 EcNAP12-92 Italy 380 LES10 2009 Greece 406 EcPAN-139A Chile 458 EcBIN-01 Chile 484 EcBAS-305 2013 Chile EcQUI-494 2013 Chile 536 EcQUI-500 2013 Chile 562 EcLAC-388 Chile 588 EcDIC-594 2013 Chile 640 EcPIL-639 2013 Chile 666 EcEST-788 Chile 692 EcEST-771 2013 Chile 116 EcQAB10-6 United Kingdom 141 EcPLY10-27 2010 United Kingdom 216 EcQB10-22 France 266 EcRIB-129 Spain 316 EcNAP12-s4-19 Italy 366 EcPAN-097A Chile 391 EcBIN-292 2013 Chile 416 EcBIN-006 Chile 441 EcCIS-071 2013 Chile 466 EcQUI-506 Chile 491 EcLAC-390 2013 Chile 17 Ec705 2006 Korea 43 OBA07-108 United Kingdom 95 EcQAB10-24 United Kingdom 121 EcREP10-50 2010 United Kingdom 173 L214 2010 France 199 EcTH10-10 France 355 EcNAP12-45 Italy 381 EcPAN-84 2013 Spain 407 EcPAN-139B Chile 459 EcBIN-02 Chile 485 EcBAS-255A 2013 Chile EcQUI-497 2013 Chile 537 EcQUI-501 2013 Chile 563 EcLAC-389 Chile 589 EcDIC-595 2013 Chile 641 EcCNA-638 2013 Chile 667 EcEST-789 Chile 693 EcEST-772 2013 Chile 117 EcQAB10-7 United Kingdom 142 EcPLY10-28 2010 United Kingdom 217 EcQB10-23 France 267 EcRIB-136 Spain 317 EcNAP12-s3-35 Italy 367 EcPAN-109 Chile 392 EcBIN-294 2013 Chile 417 EcBIN-007 Chile 442 EcPVI-035 2013 Chile 467 EcQUI-513 Chile 492 EcLAC-411 2013 Chile 18 Ec002 1988 New Zeland 44 OBA07-15 United Kingdom 96 EcQAB10-10 United Kingdom 122 EcREP10-51 2010 United Kingdom 174 L218 2010 France 200 EcTH10-12 France 356 EcNAP12-s3-51 Italy 382 EcPAN-90 2013 Spain 408 EcPAN-144A Chile 460 EcBIN-04 Chile 486 EcBAS-255D 2013 Chile EcQUI-498 2013 Chile 538 EcQUI-504 2013 Chile 564 EcLAC-390 Chile 590 EcDIC-596 2013 Chile 642 EcCNA-640 2013 Chile 668 EcEST-790 Chile 694 EcEST-775 2013 Chile 118 EcQAB10-8 United Kingdom 143 EcPLY10-29 2010 United Kingdom 218 EcQB10-24 France 268 EcGAN-516 Spain 318 EcNAP12-s3-11 Italy 368 EcPAN-110 Chile 393 EcBIN-295 2013 Chile 418 EcBIN-008 Chile 443 EcPVI-036 2013 Chile 468 EcQUI-519 Chile 493 EcLAC-415 2013 Chile 19 Ec006 1988 Australia 45 SAM08-04 United Kingdom 97 EcPLY10-19 United Kingdom 123 EcREP10-52 2010 United Kingdom 175 L219 2010 France 201 EcTH10-13 France 357 EcNAP12-s4-10 Italy 383 EcPAN-95 2013 Spain 409 EcPAN-174A Chile 461 EcBIN-05 Chile 487 EcBAS-302A 2013 Chile EcQUI-503 2013 Chile 539 EcQUI-506 2013 Chile 565 EcLAC-392 Chile 591 EcDIC-598 2013 Chile 643 EcCNA-641 2013 Chile 669 EcEST-791 Chile 695 EcEST-776 2013 Chile 119 EcQAB10-9 United Kingdom 144 EcPLY10-30 2010 United Kingdom 219 EcQB10-2 France 269 EcGAN-528 Spain 319 EcNAP12-15 Italy 369 EcPAN-113 Chile 394 EcBIN-306 2013 Chile 419 EcBIN-276 Chile 444 EcALG-520 2013 Chile 469 EcLAC-403 Chile 494 EcLAC-433 2013 Chile 20 WIC08-01 2008 United Kingdom 46 SAM08-05 United Kingdom 98 EcPLY10-20 United Kingdom 124 EcREP10-54 2010 United Kingdom 176 L220 2010 France 202 EcTH10-14 France 358 EcNAP12-s3-35 Italy 384 EcPAN-128 2013 Spain 410 EcPAN-109 Chile 462 EcBIN-06 Chile 488 EcBAS-305B 2013 Chile EcQUI-505 2013 Chile 540 EcQUI-519 2013 Chile 566 EcLAC-394 Chile 592 EcDIC-600 2013 Chile 644 EcCNA-644 2013 Chile 670 EcEST-792 Chile 696 EcEST-777 2013 Chile 120 EcQAB10-10 United Kingdom 145 EcPLY10-34 2010 United Kingdom 220 EcQB10-3 France 270 EcGAN-529 Spain 320 EcNAP12-17 Italy 370 EcPAN-114 Chile 395 EcBIN-307 2013 Chile 420 EcBIN-346 Chile 445 EcALG-521 2013 Chile 470 EcLAC-425 Chile 495 EcLAC-435 2013 Chile 21 WIC08-02 2008 United Kingdom 22 WIC08-03 2008 United Kingdom 48 SAM08-11 United Kingdom 100 EcPLY10-22 United Kingdom 126 Ec393 2003 France 178 L224 2010 France 204 EcTH10-16 France 360 EcNAP12-s3-03 Italy 386 EcPAN-140 2013 Spain 412 EcPAN-113 Chile 464 EcBIN-08 Chile 490 EcCIS-70 2013 Chile EcQUI-508 2013 Chile 542 EcQUI-450 2013 Chile 568 EcLAC-411 Chile 594 EcDIC-602 2013 Chile 646 EcCNA-660 2013 Chile 672 EcEST-794 Chile 698 EcEST-779 2013 Chile 47 SAM08-09 United Kingdom 99 EcPLY10-21 United Kingdom 125 Esil.Ros.h 1970 France 177 L222 2010 France 203 EcTH10-15 France 359 EcNAP12-s3-08 Italy 385 EcPAN-138 2013 Spain 411 EcPAN-110 Chile 463 EcBIN-07 Chile 489 EcCIS-071 2013 Chile EcQUI-507 2013 Chile 541 EcQUI-513 2013 Chile 567 EcLAC-407 Chile 593 EcDIC-601 2013 Chile 645 EcCNA-645 2013 Chile 671 EcEST-793 Chile 697 EcEST-778 2013 Chile 121 EcQAB10-11 United Kingdom 146 EcPLY10-35 2010 United Kingdom 221 EcQB10-7 France 271 EcGAN-537 Spain 321 EcNAP12-s3-13 Italy 371 EcPAN-122 Chile 396 EcBIN-308 2013 Chile 421 EcBIN-350 Chile 446 EcALG-523 2013 Chile 471 EcLAC-363 Chile 496 EcLAC-443 2013 Chile	Locality Dunstaffnage Gosport Marina Cherbourg Roscoff Roscoff Roscoff Roscoff Naples Athens, Kavouri Athens, Kavouri Athens, Agios Kosmas San Juan de Marcona Wick Wick Wick Wick Wick Wick Rattray Head Berwick Berwick Berwick Berwick Berwick Berwick Berwick Berwick Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Restronguest Restronguest Restronguest Restronguest Restronguest Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Roscoff Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Saint Malo Traezh Hir Traezh Hir Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Naples Korinthos Korinthos Korinthos Korinthos Korinthos Korinthos Korinthos Lesbos Lesbos Lesbos Lesbos Lesbos Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Puerto Viejo, Caldera Puerto Viejo, Caldera Caldera Algarrobo Algarrobo Algarrobo Algarrobo Algarrobo Algarrobo Algarrobo Algarrobo Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Dichato Dichato Dichato Dichato Dichato Dichato Cocholgüe, Tomé Cocholgüe, Tomé Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Pilolcura, Valdivia Curiñanco, Valdivia San Carlos, Corral, Valdivia San Carlos, Corral, Valdivia San Carlos, Corral, Valdivia San Carlos, Corral, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Niebla, Valdivia Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Estaquilla Pett level Pett level Pett level Pett level Pett level Pett level Pett level Pett level Pett level Pett level Pett level Gosport marina Gosport marina Gosport marina Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Plymouth Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Restronguet Roscoff Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Traezh Hir Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Ribadeo Viana do Castelo Viana do Castelo Viana do Castelo Viana do Castelo Viana do Castelo Naples Naples Naples Naples Naples Naples Naples Naples Naples Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Bahía Salada, Caldera Algarrobo Algarrobo Algarrobo Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Quintay Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Las Cruces Traezh Hir Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Las Cruces Plymouth Plymouth Traezh Hir Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Las Cruces San Juan de Marcona Berwick Plymouth Restronguest Roscoff Traezh Hir Naples Lesbos Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Pilolcura, Valdivia Estaquilla Estaquilla Plymouth Plymouth Traezh Hir Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Las Cruces Pisagua Berwick Plymouth Restronguest Saint Malo Traezh Hir Naples Lesbos Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Pilolcura, Valdivia Estaquilla Estaquilla Plymouth Plymouth Traezh Hir Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Caleta Cisne, Caldera Quintay Las Cruces Pisagua Berwick Plymouth Restronguest Saint Malo Traezh Hir Naples Lesbos Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Pilolcura, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Caleta Cisne, Caldera Quintay Las Cruces Falmouth, Massachusetts Berwick Plymouth Restronguest Saint Malo Traezh Hir Naples Lesbos Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Pilolcura, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Caleta Cisne, Caldera Quintay Las Cruces Jejudo, Kimnjung Dunstaffnage Plymouth Restronguest Saint Malo Traezh Hir Naples Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Curiñanco, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Ribadeo Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Puerto Viejo, Caldera Quintay Las Cruces Kaikoura Dunstaffnage Plymouth Restronguest Saint Malo Traezh Hir Naples Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Curiñanco, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Coruña Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Puerto Viejo, Caldera Quintay Las Cruces Victoria Dunstaffnage Plymouth Restronguest Saint Malo Traezh Hir Naples Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Curiñanco, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Coruña Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Algarrobo Las Cruces Las Cruces Wick Dunstaffnage Plymouth Restronguest Saint Malo Traezh Hir Naples Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Bahía Salada, Caldera Quintay Quintay Las Cruces Dichato Curiñanco, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Coruña Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Algarrobo Las Cruces Las Cruces Wick Wick Dunstaffnage Plymouth Roscoff Saint Malo Traezh Hir Naples Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Caleta Cisne, Caldera Quintay Quintay Las Cruces Dichato Curiñanco, Valdivia Estaquilla Estaquilla Dunstaffnage Plymouth Roscoff Saint Malo Traezh Hir Naples Pan de Azúcar Pan de Azúcar Bahía Inglesa, Caldera Caleta Cisne, Caldera Quintay Quintay Las Cruces Dichato Curiñanco, Valdivia Estaquilla Estaquilla Plymouth Plymouth Quiberon Coruña Naples Pan de Azúcar Bahía Inglesa, Caldera Bahía Inglesa, Caldera Algarrobo Las Cruces Las Cruces	Genbank Accession LM995017 LM995009 LM995014 LM995041 LM995086 LM995058 LM995005 LM995012 LM995375 LM995379 LM995368 LM995008 KT983135 KT983136 KT983137 KT983138 KT983139 KT983140 KT983384 KT983155 KT983156 KT983157 KT983158 KT983159 KT983160 KT983161 KT983162 KT982766 KT982767 KT982768 KT982769 KT982770 KT982771 KT982772 KT982773 KT982774 KT982775 KT982776 KT982777 KT982778 KT982779 KT982780 KT983096 KT983097 KT983098 KT983099 KT983100 KT983101 KT983102 KT983103 KT983104 KT983410 KT982938 KT982939 KT982940 KT982941 KT982942 KT982833 KT982834 KT983074 KT983075 KT983077 KT983078 KT983079 KT983080 KT983084 KT983085 KT983086 KT983087 KT983088 KT983089 KT983090 KT983397 KT983398 KT983399 KT983400 KT983401 KT983402 KT983403 KT983404 KT983405 KT983406 KT983409 KT982920 KT982921 KT982951 KT982952 KT982813 KT982814 KU134118 KU134119 KU134120 KU134121 KU134122 KU134123 KU134124 KU134125 KU134126 KU134127 KU134128 KU134141 KU134142 KU134154 KU134155 KU134156 KU134129 KU134130 KU134131 KU134132 KU134133 KU134134 KU134135 KT982913 KT982914 KU134116 KU134117 KU134136 KT982874 KT982875 KT982876 KT982966 KT982967 KT982968 KT982969 KT982970 KT982971 KT982972 KT983194 KT983195 KT983196 KT983197 KT983198 KT982890 KT982891 KT982892 KT982893 KT982894 KT982895 KT982896 KT982897 KT982898 KT982899 KT982900 KT982901 KT982902 KT982903 KT982904 KU134022 KU134024 KU134025 KU134026 KU134027 KU134028 KU134029 KU134030 KU134032 KU134037 KU134038 KU134039 KU134040 KT982907 KU134031 KU134023 KT987930 KU134105 KT982995 KT982996 KT982997 KT982998 KT982999 KT983000 KT983001 KU134108 KT982974 KT982975 KT982976 KT982977 KT982989 KT982990 KT982991 KT982992 KT982993 KT982994 KT987960 KT987961 KT987962 KT987963 KT987964 KT987965 KT987966 KT987967 KT987968 KU134054 KU134055 KU134056 KU134106 KU134107 KT987932 KT987933 KT987934 KT987935 KT987936 KT987937 KT987938 KT987939 KT987940 KT987941 KT987953 KT987954 KT987955 KT987956 KT987957 KT987958 KT987959 KT987974 KT987977 KT983009 KU134069 KU134070 KU134071 KU134072 KU134073 KT987970 KT987975 KT983010 KT983011 KT983012 KT983013 KT983014 KT983015 KT983016 KT983224 KT983225 KT983226 KT983227 KT983228 KT983231 KT983229 KT983230 KU134063 KU134064 KU134065 KU134057 KU134096 KU134097 KU134098 KU134099 KU134113 KU134114 KU134115 KT987972 KT987973 KT983039 KT983040 KT983041 KT983042 KT983043 KT983044 KT983045 KT983046 KT983047 KT983048 KT983049 KT983050 KT983051 KT983052 KT983053 KU134161 KU134305 KU134306 KU134307 KU134308 KU134309 KU134310 KU134311 KU134312 KU134313 KU134314 KU134542 KU134543 KU134544 KU134213 KU134224 KU134225 KU134226 KU134227 KU134228 KU134229 KU134230 KU134231 KU134232 KU134233 KU134234 KU134235 KU134236 KU134472 KU134473 KU134540 KU134315 KU134316 KU134317 KU134318 KU134319 KU134320 KU134321 KU134322 KU134323 KU134324 KU134325 KU134326 KU134470 KU134333 KU134334 KU134335 KU134336 KU134337 KU134338 KU134339 KU134340 KU134341 KU134408 KU134409 KU134410 KU134411 KU134412 KU134293 KU134294 KU134295 KU134296 KU134297 KU134298 KU134299 KU134300 KU134301 KU134302 KU134484 KU134485 KU134486 KU134487 KU134343 KU134344 KU134508 KU134509 KU134510 KU134252 KU134253 KU134254 KU134255 KU134256 KU134257 KU134258 KU134259 KU134260 KU134172 KU134173 KU134174 KU134175 KU134176 KU134177 KU134178 KU134640 KU134641 KU134642 KU134643 KU134644 KU134645 KU134646 KU134709 KU134710 KU134711 KU134712 KU134713 KU134714 KU134715 KU134716 KU134717 KU134179 KU134180 KU134181 KU134182 KU134183 KU134195 KU134197 KU134198 KU134199 KU134200 KU134201 KU134202 KU134203 KU134204 KU134205 KU134206 KU134207 KU134595 KU134596 KU134616 KU134617 KU134618 KU134619 KU134620 KU134621 KU134622 KU134593 KU134600 KU134601 KU134602 KU134603 KU134604 KU134598 KU134350 KU134351 KU134677 KU134550 KU134554 KU134555 KU134556 KU134557 KU134563 KU134570 KU134571 KU134573 KU134629 KU134718 KU134551 KU134552 KU134553 KU134559 KU134560 KU134561 KU134562 KU134566 KU134567 KU134568 KU134569 KU134574 KU134575 KU134576 KU134413 KU134488 KU134691 KU134698 KU134184 KU134597 KU134599 KU134719 KU134577 KU134214 KU134474 KU134414 KU134489 KU134692 KU134699 KU134185 KU134605 KU134606 KU134720 KU134578 FP885846 KT983163 KT982781 KT982943 KT982815 KT982953 KU134143 KU134137 KT983199 KT982905 KU134033 KT982978 KT983214 KT987942 KU134074 KT983232 KT983028 KT983054 KU134215 KU134475 KU134415 KU134490 KU134693 KU134700 KU134186 KU134607 KU134196 KU134721 KU134579 LM995007 KT983164 KT982782 KT982944 KT982916 KT982954 KU134144 KU134138 KT983200 KT982906 KU134034 KT982979 KT983215 KT987943 KU134075 KT983233 KT983029 KT983055 KU134216 KU134476 KU134416 KU134491 KU134694 KU134701 KU134187 KU134608 KU134548 KU134722 KU134580 LM995010 KT983165 KT982783 KT982945 KT982917 KT982955 KU134145 KU134139 KT983201 KU134013 KU134035 KT982980 KT983216 KT987944 KU134076 KU134090 KT983030 KT983237 KU134217 KU134477 KU134237 KU134492 KU134695 KU134702 KU134188 KU134609 KU134624 KU134723 KU134581 LM995016 KT983166 KT982784 KT982946 KT982918 KT982956 KU134146 KU134140 KT983202 KU134014 KU134036 KT982981 KT983217 KT987945 KU134077 KT987971 KT983031 KT983238 KU134218 KU134478 KU134238 KU134493 KU134696 KU134703 KU134189 KU134610 KU134625 KU134724 KU134582 LM995011 KT982922 KT982785 KT982947 KT982919 KT982957 KU134147 KT982867 KT983203 KU134015 KU134041 KT982982 KT983218 KT987946 KU134078 KT983017 KT983032 KT983239 KU134219 KU134479 KU134239 KU134494 KU134697 KU134704 KU134190 KU134611 KU134565 KU134725 KU134583 LM995018 KT982923 KT982786 KT982948 KT983388 KT982958 KU134148 KT982868 KT983204 KU134016 KU134042 KT982983 KT983219 KT987947 KU134079 KT983018 KT983033 KT983240 KU134220 KU134480 KU134240 KU134497 KU134727 KU134705 KU134191 KU134612 KU134594 KU134726 KU134584 LM995019 KT982924 KT983091 KT982949 KT983389 KT982959 KU134149 KT982869 KT983205 KU134017 KU134043 KT982984 KT983220 KT987948 KU134080 KT983019 KT983034 KT983241 KU134221 KU134481 KU134241 KU134278 KU134729 KU134706 KU134192 KU134613 KU134347 KU134346 KU134585 KT983131 KT982925 KT983092 KT982950 KT983390 KT982960 KU134150 KT982870 KT983206 KU134018 KU134044 KT982985 KT983221 KT987949 KU134081 KT983020 KT983035 KT983242 KU134222 KU134482 KU134242 KU134279 KU134730 KU134707 KU134193 KU134614 KU134348 KU134546 KU134587 KT983132 KT983133 KT983383 KT983094 KT982817 KT983392 KT982962 KU134152 KT982872 KT983208 KU134020 KU134053 KT982987 KU134052 KT987951 KU134083 KT983022 KT983037 KT983244 KT982926 KT983093 KT982830 KT983391 KT982961 KU134151 KT982871 KT983207 KU134019 KU134051 KT982986 KT983222 KT987950 KU134082 KT983021 KT983036 KT983243 KU134223 KU134483 KU134243 KU134280 KU134731 KU134708 KU134194 KU134615 KU134349 KU134547 KU134588	Reference Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper Peters et al. 2015 This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper
						13

Table S3 .

 S3 Mitochondrial (COI-5P) and nuclear (ITS1) DNA sequence variation in E. siliculosus. Molecular diversity indices were 1 calculated for the two molecular markers (COI-5P and ITS1). N= number of sequences; nH= number of haplotypes; H= gene diversity; 2

	3	π= nucleotide diversity; S= number of polymorphic sites. Standard deviations (SD) in parentheses.
						COI-5P			ITS1
		Region	Site	Code N nH	H (SD)	π (SD) (.10 -2 ) S	N nH	H (SD)	π (SD) (.10 -2 ) S
		United	Dunstaffnage	DUN 4	2	0.500 (0.265) 0.166 (0.164) 2	4	3	0.833 (0.222) 0.343 (0.279) 4
		Kingdom	Pett level	PET 10 3	0.378 (0.181) 0.066 (0.076) 2	10 4	0.778 (0.091) 0.450 (0.288) 7
			Plymouth	PLY 23 2	0.403 (0.091) 0.067 (0.072) 1	23 6	0.739 (0.064) 0.344 (0.217) 8
			Restronguet	RES 12 3	0.591 (0.108) 0.116 (0.106) 2	12 6	0.849 (0.074) 0.333 (0.221) 9
			Total UK	49 5	0.501 (0.060) 0.102 (0.091) 5	49 12 0.850 (0.029) 0.468 (0.273) 13
		France	Roscoff	ROS 16 2	0.125 (0.106) 0.021 (0.038) 1	16 10 0.892 (0.063) 0.445 (0.274) 13
			Traezh Hir	THZ 15 4	0.619 (0.120) 0.221 (0.163) 4	15 4	0.619 (0.120) 0.438 (0.271) 11
			Quiberon	QUI 14 2	0.363 (0.130) 0.060 (0.069) 1	14 4	0.648 (0.116) 0.344 (0.224) 6
			Total France	45 5	0.

549 (0.065) 0.123 (0.103) 5 45 16 0.848 (0.040) 0.534 (0.305) 19

  TableS4. Mitochondrial (COI-5P) and nuclear (ITS1) DNA sequence variation in E. crouaniorum. Molecular diversity indices were 1 calculated for the two molecular markers (COI-5P and ITS1). N= number of sequences; nH= number of haplotypes; H= gene diversity; 2 π= nucleotide diversity; S= number of polymorphic sites. Standard deviations (SD) in parentheses. 3

						COI-5P					ITS1
	Region	Site	Code N nH	H (SD)	π (SD) (.10 -2 ) S	N nH		H (SD)	π (SD) (.10 -2 ) S
	United	Wick	WIC	8	1	0	0	0	8	7	0.964 (0.077) 0.846 (0.507) 19
	Kingdom Berwick	BER 11	2	0.436 (0.133) 0.072 (0.079) 1	11 5	0.855 (0.066) 0.234 (0.162)	5
		Mull of Galloway MUL 12	3	0.318 (0.164) 0.055 (0.067) 2	12 9	0.939 (0.058) 0.626 (0.367) 20
		Plymouth	PLY 11	3	0.618 (0.104) 0.302 (0.212) 4	11 7	0.891 (0.074) 0.946 (0.538) 20
		Total UK	42	6	0.474 (0.087) 0.164 (0.126) 6	42 23 0.938 (0.024) 0.728 (0.393) 36
	France	Roscoff	ROS 15	5	0.562 (0.143) 0.190 (0.145) 6	15 12 0.962 (0.040) 0.663 (0.379) 28
		Traezh Hir	THZ	9	3	0.556 (0.165) 0.101 (0.100) 2	9	7	0.917 (0.092) 0.493 (0.308) 16
		Quiberon	QUI 12	6	0.758 (0.122) 0.183 (0.145) 5	12 6	0.849 (0.074) 0.269 (0.176)	8
		Total France	36 10 0
	NW Iberia	Ribadeo	RIB 13 3	0.615 (0.078) 0.140 (0.119) 3	13 4	0.692 (0.115) 0.314 (0.209) 7
		Coruña	COR 13 3	0.295 (0.156) 0.051 (0.063) 2	13 8	0.808 (0.113) 0.321 (0.213) 12
		Viana	VIA	4	2	0.500 (0.265) 0.166 (0.164) 2	4		4	1.000 (0.177) 0.293 (0.246) 4
		Total Iberia	30 6	0.529 (0.095) 0.121 (0.103) 7	30 13 0.862 (0.045) 0.338 (0.211) 18
	Mediterranean Naples	NAP	9	2	0.389 (0.164) 0.065 (0.075) 1	9		2	0.389 (0.164) 0.057 (0.066) 1
	Chile	Pan de Azúcar	PAN 10 2	0.356 (0.159) 0.118 (0.109) 2	10 2	0.200 (0.154) 0.029 (0.045) 1
		Caldera	CAL 29 5	0.665 (0.067) 0.169 (0.130) 5	29 10 0.796 (0.062) 0.647 (0.366) 16
		Concepción	CON	1	1	0	0	0	1		1	0	0
		Achao	ACH	4	1	0	0	0	4		4	1.00 (0.177)	0.606 (0.453) 7

.624 (0.092) 0.170 (0.130) 11 36 22 0.916 (0.037) 0.523 (0.295) 38

	NW	Ribadeo		8	1	0	0	0	8	5	0.893 (0.086) 0.694 (0.423) 14
	Iberia	Coruña	RIB	5	3	0.800 (0.164) 0.166 (0.154) 2	5	5	1.000 (0.127) 0.753 (0.504) 15
		Ría de Arousa	COR	5	2	0.600 (0.175) 0.100 (0.109) 1	5	4	0.900 (0.161) 0.243 (0.191)	4
		Viana	VIA	2	2	1.000 (0.500) 0.332 (0.406) 2	2	2	1.000 (0.500) 1.453 (1.512) 12
		Total Iberia		20	5	0.679 (0.080) 0.148 (0.121) 4	20 14 0.963 (0.026) 0.711 (0.396) 40
	Chile	Las Cruces	LAC	6	3	0.600 (0.215) 0.111 (0.112) 2	6	4	0.800 (0.172) 0.543 (0.359) 11
		Concepción	CON	7	2	0.571 (0.120) 0.095 (0.099) 1	7	5	0.905 (0.103) 0.984 (0.596) 16
		Valdivia	VAL 11	2	0.327 (0.153) 0.054 (0.066) 1	11 6	0.800 (0.114) 0.466 (0.286) 18
		Estaquilla	EST 24	1	0	0	0	24 8	0.808 (0.053) 0.668 (0.371) 14
		Achao	ACH	4	1	0	0	0	4	4	1.000 (0.177) 0.364 (0.285)	5
		Total Chile		52	3	0.429 (0.059) 0.073 (0.073) 2	52 23 0.928 (0.019) 0.836 (0.

444) 31 TOTAL 150 18 0.583 (0.046) 0.154 (0.118) 18 150 68 0.953 (0.010) 0.759 (0.401) 80Table S5 .

 S5 Mitochondrial (COI-5P) and nuclear (ITS1) DNA sequence variation in Ectocarpus 6. Molecular diversity indices were calculated for the two molecular markers (COI-5P and ITS1). N= number of sequences; nH= number of haplotypes; H= gene diversity; π= nucleotide diversity; S= number of polymorphic sites. Standard deviations (SD) in parentheses.

				COI-5P					ITS1
	Site	Code N nH	H (SD)	π (SD) (.10 -2 )	S	N	nH	H (SD)	π (SD) (.10 -2 )	S
	Pisagua	PIS	1 1	0	0	0	1	1			0
	Caldera	CAL	2 2	1.000 (0.500)	0.829 (0.908)	5	2	2	1.000 (0.500)	0.267 (0.377)	1
	Quintay	QUI	9 4	0.750 (0.112)	0.525 (0.340)	7	9	5	0.806 (0.120)	0.781 (0.511)	11
	Las Cruces	LAC 29 6	0.791 (0.048)	0.440 (0.269)	8	29	19	0.934 (0.034)	1.953 (1.046)	38
	Concepción	CON	2 1	0	0	0	2	2	1.000 (0.500)	0.533 (0.653)	2
	Valdivia	VAL	3 3	1.000 (0.272)	0.663 (0.564)	6	3	2	0.667 (0.314)	2.667 (2.104)	15
	TOTAL		46 8	0.822 (0.022)	0.511 (0.300) 10	46	25	0.897 (0.037)	1.615 (0.868)	45

Table S6 .

 S6 Species nomenclature used in literature for the species of the Ectocarpus siliculosi group. 1

Montecinos et al. Stache-Crain et al. 1997 Peters et al. 2010a Peters et al. 2010b Peters et al. 2015

	E. siliculosus	1a	E. siliculosus	GT6-1a	E. siliculosus
	E. crouaniorum	2c	E. crouaniorum	GT8-2c	E. crouaniorum
	Ectocarpus 1	4	E. siliculosus (?)	4	4
	Ectocarpus 2	4	---	---	4
	Ectocarpus 3	3	---	---	3
	Ectocarpus 4	---	---	---	---
	Ectocarpus 5	---	---	---	1d
	Ectocarpus 6	---	---	GT1	1c
	Ectocarpus 7	1c	E. siliculosus	GT4-1c	1c
	Ectocarpus 8	1b	---	---	---
	Ectocarpus 9	1b	---	---	---
	Ectocarpus 10	1b	---	GT2-1b	1b
	Ectocarpus 11	---	---	GT3	1b
	Ectocarpus 12	2b	---	GT8-2b	2b
	Ectocarpus 13	2a	---	---	2a

Journal of Phycology

Acknowledgments

We are particularly grateful to C. Destombe, S. Faugeron, A. Mann, D.G. Müller, P. Murúa, D. Patiño, D. Schroeder, J. West and R. Westermeier, who helped by collecting samples. The authors thank Biogenouest Core Facility for technical assistance and the members of Grupo BioCost of Coruña University for advice and logistic support during sampling in Galicia and North Portugal. We also thank to S. Mauger for assistance in the molecular laboratory and L. Dartevelle and J. Coudret for help in maintaining strains in the Station Biologique de Roscoff. We particularly appreciated insightful and constructive comments from C. Destombe and J. M. Cock. Principal funding came from

GWY08-05 2008 United Kingdom Mull of Galloway KT983141 This paper 50 GWY08-07 2008 United Kingdom Mull of Galloway KT983142 This paper 51 GWY08-08 2008 United Kingdom Mull of Galloway KT983143 This paper 52 GWY08-09 2008 United Kingdom Mull of Galloway KT983144 This paper 53 GWY08-13 2008 United Kingdom Mull of Galloway KT983145 This paper 54 GWY08-14 2008 United Kingdom Mull of Galloway KT983146 This paper 55 GWY08-15 2008 United Kingdom Mull of Galloway KT983147 This paper 56 GWY08-16 2008 United Kingdom Mull of Galloway KT983148 This paper 57 GWY08-17 2008 United Kingdom Mull of Galloway KT983149 This paper 58 GWY08-21 2008 United Kingdom Mull of Galloway KT983150 This paper 59 GWY08-22 2008 United Kingdom Mull of Galloway KT983151 This paper 60 GWY08-23 2008 United Kingdom Mull of Galloway KT983152 This paper 61 GWY08-24 2008 United Kingdom Mull of Galloway KT983153 This paper 62 GWY08-26 2008 United Kingdom Mull of Galloway KT983154 This paper 63 HAS08-06 2008 United Kingdom Pett level KT982927 This paper 64 HAS08-07 2008 United Kingdom Pett level KT982928 This paper 65 HAS08-09 2008 United Kingdom Pett level KT982929 This paper 66 HAS08-12 2008 United Kingdom Pett level KT982930 This paper 67 HAS08-13 2008 United Kingdom Pett level KT982931 This paper 68 HAS08-14 2008 United Kingdom Pett level KT982932 This paper 69 HAS08-16 2008 United Kingdom Pett level KT982933 This paper 70 HAS08-17 2008 United Kingdom Pett level KT982934 This paper 71 HAS08-18 2008 United Kingdom Pett level KT982935 This paper 72 HAS08-19 2008 United Kingdom Pett level KT982936 This paper 73 HAS08-20 2008 United Kingdom Pett level KT982937 This paper 74 EcQAB10-001 2010 United Kingdom Plymouth KT982764 This paper France Roscoff KT983083 This paper 140 EcPH11-s5-13 France Roscoff KT982816 This paper 141 EcPH11-s5-14 France Roscoff KT982818 This paper 142 EcPH11-s5-15 France Roscoff KT982819 This paper 143 EcPH11-s5-17 France Roscoff KT982820 This paper 144 EcPH11-s5-18 France Roscoff KT982821 This paper 145 EcPH11-s5-19 France Roscoff KT982822 This paper 146 EcPH11-s5-20 France Roscoff KT982823 This paper 147 EcPH11-s2B-41 France Roscoff KT982824 This paper 148 EcPH11-s2B-43 France Roscoff KT982825 This paper 149 EcPH11-s2B-45 France Roscoff KT982826 This paper 150 EcPH11-s-2A-06 France Roscoff KT982827 This paper 151 EcPH11-s-2A-04 France Roscoff KT982828 This paper 152 BLZ11-229 France Roscoff KT982829 This paper France Roscoff KU134456 This paper 162 EcPH10-38 France Roscoff KU134455 This paper 163 EcPH10-72 France Roscoff KU134471 This paper 164 EcPH11-s5-15 France Roscoff KU134261 This paper 165 EcPH11-s2B-45 France Roscoff KU134262 This paper 166 EcPH11-s5-13 France Roscoff KU134263 This paper 167 EcPH11-s2B-43 France Roscoff KU134264 This paper 168 EcPH11-s5-19 France Roscoff KU134265 This paper 169 EcPH11-s2B-41 France Roscoff KU134266 This paper 170 EcPH11-s5-18 France Roscoff KU134267 This paper 171 EcPH11-s5-17 France Roscoff KU134268 This paper