Seagrasses colonized the sea 1 on at least three independent occasions to form the basis of one of the most productive and widespread coastal ecosystems on the planet 2 . Here we report the genome of Zostera marina (L.), the first, to our knowledge, marine angiosperm to be fully sequenced. This reveals unique insights into the genomic losses and gains involved in achieving the structural and physiological adaptations required for its marine lifestyle, arguably the most severe habitat shift ever accomplished by flowering plants. Key angiosperm innovations that were lost include the entire repertoire of stomatal genes 3 , genes involved in the synthesis of terpenoids and ethylene signalling, and genes for ultraviolet protection and phytochromes for far-red sensing. Seagrasses have also regained functions enabling them to adjust to full salinity. Their cell walls contain all of the polysaccharides typical of land plants, but also contain polyanionic, low-methylated pectins and sulfated galactans, a feature shared with the cell walls of all macroalgae 4 and that is important for ion homoeostasis, nutrient uptake and O 2 /CO 2 exchange through leaf epidermal cells. The Z. marina genome resource will markedly advance a wide range of functional ecological studies from adaptation of marine ecosystems under climate warming 5,6 , to unravelling the mechanisms of osmoregulation under high salinities that may further inform our understanding of the evolution of salt tolerance in crop plants 7
We review the progress made in the emerging field of coastal seascape ecology, i.e. the application of landscape ecology concepts and techniques to the coastal marine environment. Since the early 1990s, the landscape ecology approach has been applied in several coastal subtidal and intertidal biogenic habitats across a range of spatial scales. Emerging evidence indicates that animals in these seascapes respond to the structure of patches and patch mosaics in different ways and at different spatial scales, yet we still know very little about the ecological significance of these relationships and the consequences of change in seascape patterning for ecosystem functioning and overall biodiversity. Ecological interactions that occur within patches and among different types of patches (or seascapes) are likely to be critically important in maintaining primary and secondary production, trophic transfer, biodiversity, coastal protection, and supporting a wealth of ecosystem goods and services. We review faunal responses to patch and seascape structure, including effects of fragmentation on 5 focal habitats: seagrass meadows, salt marshes, coral reefs, mangrove forests, and oyster reefs. Extrapolating and generalizing spatial relationships between ecological patterns and processes across scales remains a significant challenge, and we show that there are major gaps in our understanding of these relationships. Filling these gaps will be crucial for managing and responding to an inevitably changing coastal environment. We show that critical ecological thresholds exist in the structural patterning of biogenic ecosystems that, when exceeded, cause abrupt shifts in the distribution and abundance of organisms. A better understanding of faunal-seascape relationships, including the identifications of threshold effects, is urgently needed to support the development of more effective and holistic management actions in restoration, site prioritization, and forecasting the impacts of environmental change.
As the most widespread seagrass in temperate waters of the Northern Hemisphere, Zostera marina provides a unique opportunity to investigate the extent to which the historical legacy of the last glacial maximum (LGM18 000-10 000 years bp) is detectable in modern population genetic structure. We used sequences from the nuclear rDNA-internal transcribed spacer (ITS) and chloroplast matK-intron, and nine microsatellite loci to survey 49 populations (> 2000 individuals) from throughout the species' range. Minimal sequence variation between Pacific and Atlantic populations combined with biogeographical groupings derived from the microsatellite data, suggest that the trans-Arctic connection is currently open. The east Pacific and west Atlantic are more connected than either is to the east Atlantic. Allelic richness was almost two-fold higher in the Pacific. Populations from putative Atlantic refugia now represent the southern edges of the distribution and are not genetically diverse. Unexpectedly, the highest allelic diversity was observed in the North Sea-Wadden Sea-southwest Baltic region. Except for the Mediterranean and Black Seas, significant isolation-by-distance was found from ~150 to 5000 km. A transition from weak to strong isolation-by-distance occurred at ~150 km among northern European populations suggesting this scale as the natural limit for dispersal within the metapopulation. Links between historical and contemporary processes are discussed in terms of the projected effects of climate change on coastal marine plants. The identification of a high genetic diversity hotspot in Northern Europe provides a basis for restoration decisions.
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