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  • Genetic diversity of Saccharina latissima (Phaeophyceae) along a salinity gradient in the North Sea-Baltic Sea transition zone
    Publication . Nielsen, Mette Moller; Paulino, Cristina; Neiva, J.; Krause-Jensen, Dorte; Bruhn, Annette; Serrão, Ester
    The North Sea-Baltic Sea transition zone constitutes a boundary area for the kelp species Saccharina latissima due to a strong salinity gradient operating in the area. Furthermore, the existence of S.latissima there, along Danish waters, is fairly patchy as hard bottom is scarce. In this study, patterns of genetic diversity of S.latissima populations were evaluated along the salinity gradient area of Danish waters (here designated brackish) and were compared to reference sites (here designated marine) outside the gradient area, using microsatellite markers. The results showed that the S.latissima populations were structured into two clusters corresponding to brackish versus marine sites, and that gene flow was reduced both between clusters and between populations within clusters. In addition, results provided empirical evidence that marginal populations of S.latissima in the salinity gradient area exhibited a distinct genetic structure when compared to marine ones. Brackish populations were less diverse, more related, and showed increased differentiation over distance compared to marine populations. The isolation of the brackish S.latissima populations within the salinity gradient area of Danish waters in conjunction with their general low genetic diversity makes these populations vulnerable to ongoing environmental and climate change, predicted to result in declining salinity in the Baltic Sea area that may alter the future distribution and performance of S.latissima in the area.
  • Genetic structure of amphi-Atlantic Laminaria digitata (Laminariales, Phaeophyceae) reveals a unique range-edge gene pool and suggests post-glacial colonization of the NW Atlantic
    Publication . Neiva, J.; Serrao, Ester; Paulino, Cristina; Gouveia, Licínia; Want, Andrew; Tamigneaux, Éric; Ballenghien, Marion; Mauger, Stéphane; Fouqueau, Louise; Engel-Gautier, Carolyn; Destombe, Christophe; Valero, Myriam
    In the North-east (NE) Atlantic, most intertidal fucoids and warm-temperate kelps show unique low-latitude gene pools matching long-term climatic refugia. For cold-temperate kelps data are scarcer despite their unique cultural, ecological and economic significance. Here we test whether the amphi-Atlantic range of Laminaria digitata is derived from past glacial survival (and vicariance) in both NE and North-west (NW) Atlantic refugia (as suggested by niche modelling), or post-glacial (re)colonization (as suggested by low mtDNA divergence). We screened 14 populations from across the species range for 12 microsatellite loci to identify and map major gene pools and refugia. We assessed if NW Atlantic survival was supported by unique endemic variation, and if genetic diversity and structure were, as predicted from larger hindcasted glacial ranges, higher in the NE Atlantic. Microsatellite data subdivided L. digitata into three main genetic groups matching Brittany, northern Europe and the NW Atlantic, with finer-scale sub-structuring within European clusters. The relatively diverse NE Atlantic lineages probably survived the Last Glacial Maximum along unglaciated periglacial shorelines of the Armorican and Celtic Seas (Brittany cluster) and Ireland (northern European cluster), and remain well differentiated despite their relative proximity. The unique Brittany gene pool, at the contemporary European rear edge, is projected to disappear in the near future under high greenhouse gas emission scenarios. Low allelic diversity and low endemism in the NW Atlantic are consistent with recent post-glacial colonization from Europe, challenging the long-standing hypothesis of in situ glacial survival. Confusion with Hedophyllum nigripes may have led to underestimation of regional diversity of L. digitata, but also to overestimation of its presence along putative trans-Atlantic migration routes. Partial incongruence between modelling and genetic-based biogeographic inferences highlights the benefits of comparing both approaches to understand how shifting climatic conditions affect marine species distributions and explain large-scale patterns of spatial genetic structure.
  • Ecological traits, genetic diversity and regional distribution of the macroalga Treptacantha elegans along the Catalan coast (NW Mediterranean Sea)
    Publication . Medrano, Alba; Hereu, Bernat; Mariani, Simone; Neiva, J.; Pagès-Escolà, Marta; Paulino, Cristina; Rovira, Graciel·la; Serrao, Ester; Linares, Cristina
    The widespread decline of canopy-forming macroalgal assemblages has been documented in many regions during the last decades. This pattern is often followed by the replacement of structurally complex algal canopies by more simplified habitats (e.g., turfs or sea urchin barren grounds). Against all odds, the fucoid Treptacantha elegans, a large Mediterranean brown macroalga, broadened its depth range to deeper and exposed environments and displayed an unexpected range expansion along the northern coast of Catalonia over the last two decades. Here, we reconstruct the spread of T. elegans in time and space and unravel ecological and demographic traits such as population dynamics and genetic patterns to provide a comprehensive and integrated view of the current status and geographical expansion for this species. Fast-growing dynamics, early fertile maturity, and high turnover rate are the main competitive advantages that allow the exposed populations of T. elegans to colonize available substrata and maintain dense and patchy populations. We also provided evidence that the deeper and exposed populations of T. elegans constitute a single group across the Catalan coast, with little genetic differentiation among populations. This seems to support the hypothesis of a unique source of spread in the last decades from the Medes Islands No-Take Zone towards both southern and northern waters.
  • Cryptic diversity, geographical endemism and allopolyploidy in NE Pacific seaweeds
    Publication . Neiva, J.; Serrão, Ester; Anderson, Laura; Raimondi, Peter T.; Martins, Neusa; Gouveia, Licínia; Paulino, Cristina; Coelho, Nelson C.; Miller, Kathy A.; Reed, Daniel C.; Ladah, Lydia; Pearson, G. A.
    Background Molecular markers are revealing a much more diverse and evolutionarily complex picture of marine biodiversity than previously anticipated. Cryptic and/or endemic marine species are continually being found throughout the world oceans, predominantly in inconspicuous tropical groups but also in larger, canopy-forming taxa from well studied temperate regions. Interspecific hybridization has also been found to be prevalent in many marine groups, for instance within dense congeneric assemblages, with introgressive gene-flow being the most common outcome. Here, using a congeneric phylogeographic approach, we investigated two monotypic and geographically complementary sister genera of north-east Pacific intertidal seaweeds (Hesperophycus and Pelvetiopsis), for which preliminary molecular tests revealed unexpected conflicts consistent with unrecognized cryptic diversity and hybridization. Results The three recovered mtDNA clades did not match a priori species delimitations. H. californicus was congruent, whereas widespread P. limitata encompassed two additional narrow-endemic species from California - P. arborescens (here genetically confirmed) and P. hybrida sp. nov. The congruence between the genotypic clusters and the mtDNA clades was absolute. Fixed heterozygosity was apparent in a high proportion of loci in P. limitata and P. hybrida, with genetic analyses showing that the latter was composed of both H. californicus and P. arborescens genomes. All four inferred species could be distinguished based on their general morphology. Conclusions This study confirmed additional diversity and reticulation within NE Pacific Hesperophycus/Pelvetiopsis, including the validity of the much endangered, modern climatic relict P. arborescens, and the identification of a new, stable allopolyploid species (P. hybrida) with clearly discernable ancestry (♀ H. californicus x ♂ P. arborescens), morphology, and geographical distribution. Allopolyploid speciation is otherwise completely unknown in brown seaweeds, and its unique occurrence within this genus (P. limitata possibly representing a second example) remains enigmatic. The taxonomic separation of Hesperophycus and Pelvetiopsis is not supported and the genera should be synonymized; we retain only the latter. The transitional coastline between Point Conception and Monterey Bay represented a diversity hotspot for the genus and the likely sites of extraordinary evolutionary events of allopolyploid speciation at sympatric range contact zones. This study pinpoints how much diversity (and evolutionary processes) potentially remains undiscovered even on a conspicuous seaweed genus from the well-studied Californian intertidal shores let alone in other, less studied marine groups and regions/depths.
  • Characterization of 12 polymorphic microsatellite markers in the sugar kelp Saccharina latissima
    Publication . Paulino, Cristina; Neiva, J.; Coelho, Nelson C.; Aires, Tânia; Marba, Núria; Krause-Jensen, Dorte; Serrão, Ester
    Saccharina latissima is an ecologically and economically important kelp species native to the coastal regions of the Northern Hemisphere. This species has considerable phylogeographic structure and morphological plasticity, but lack of resolution of available genetic markers prevents a finer characterization of its genetic diversity and structure. Here, we describe 12 microsatellite loci identified in silico in a genomic library, and assess their polymorphism in three distant populations. Allelic richness at the species level was relatively high (5-23 alleles per locus), as was gene diversity within populations (0.42 < H (E) < 0.62). In addition, individuals readily form distinct genotypic clusters matching their populations of origin. The variation detected confirms the great potential of these markers to investigate the biogeography and population dynamics of S. latissima, and to better characterize its genetic resources for the establishing farming industry.
  • Trans-Arctic asymmetries, melting pots and weak species cohesion in the low-dispersal amphiboreal seaweed Fucus distichus
    Publication . Neiva, J.; Assis, Jorge; Fragkopoulou, Eliza; Pearson, Gareth; Raimondi, Peter T.; Anderson, Laura; Krause-Jensen, Dorte; Marbà, Núria; Want, Andrew; Selivanova, Olga; Nakaoka, Masahiro; Grant, W. Stewart; Konar, Brenda; Roleda, Michael Y.; Sejr, Mikael K.; Paulino, Cristina; Serrao, Ester
    Amphiboreal taxa are often composed of vicariant phylogroups and species complexes whose divergence and phylogeographic affinities reflect a shared history of chronic isolation and episodic trans-Arctic dispersal. Ecological filters and shifting selective pressures may also promote selective sweeps, niche shifts and ecological speciation during colonization, but these are seldom considered at biogeographical scales. Here we integrate genetic data and Ecologic Niche Models (ENMs) to investigate the historical biogeography and cohesion of the polymorphic rockweed Fucus distichus throughout its immense amphiboreal range, focusing on trans-Arctic asymmetries, glacial/interglacial dynamics, and integrity of sympatric eco-morphotypes. Populations were sampled throughout the Pacific and the Atlantic, from southern rear-edges to the high-Arctic. They were genotyped for seven microsatellites and an mtDNA spacer, and genetic diversity and structure were assessed from global to local scales. ENMs were used to compare niche divergence and magnitude of post-glacial range shifts in Pacific versus Atlantic sub-ranges. Haplotypic and genotypic data revealed distinct and seemingly isolated Pacific vs Arctic/Atlantic gene-pools, with finer-scale regional sub-structuring pervasive in the Pacific. MtDNA diversity was highly structured and overwhelmingly concentrated in the Pacific. Regionally, Alaska showed the highest intra-population diversity but the lowest levels of endemism. Some sympatric/parapatric ecotypes exhibited distinct genotypic/haplotypic compositions. Strikingly, niche models revealed higher Pacific tolerance to maximum temperatures and predicted a much more consolidated presence in the NE Atlantic. Glacial and modern ranges overlapped extensively in the Pacific, whereas the modern Atlantic range was largely glaciated or emerged during the Last Glacial Maximum. Higher genetic and ecogeographic diversity supports a primary Pacific diversification and secondary Atlantic colonization, also likely reflecting the much larger and more stable climatic refugia in the Pacific. The relic distribution and reduced ecological/morphological plasticity in the NE Atlantic are hypothesized to reflect functional trans-Arctic bottlenecks, recent colonization or competition with congeners. Within the Pacific, Alaska showed signatures of a post-glacial melting pot of eastern and southern populations. Genetic/ecotypic variation was generally not sufficiently discontinuous or consistent to justify recognizing multiple taxonomic entities, but support a separate species in the eastern Pacific, at the southern rear-edge. We predict that layered patterns of phylogeographic structure, incipient speciation and niche differences might be common among widespread low-dispersal amphiboreal taxa.