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

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BackgroundSignificant effects of recent global climate change have already been observed in a variety of ecosystems, with evidence for shifts in species ranges, but rarely have such consequences been related to the changes in the species genetic pool. The stretch of Atlantic coast between North Africa and North Iberia is ideal for studying the relationship between species distribution and climate change as it includes the distributional limits of a considerable number of both cold- and warm-water species.We compared temporal changes in distribution of the canopy-forming alga Fucus vesiculosus with historical sea surface temperature (SST) patterns to draw links between range shifts and contemporary climate change. Moreover, we genetically characterized with microsatellite markers previously sampled extinct and extant populations in order to estimate resulting cryptic genetic erosion.ResultsOver the past 30 years, a geographic contraction of the southern range edge of this species has occurred, with a northward latitudinal shift of approximately 1,250 km. Additionally, a more restricted distributional decline was recorded in the Bay of Biscay. Coastal SST warming data over the last three decades revealed a significant increase in temperature along most of the studied coastline, averaging 0.214°C/decade. Importantly, the analysis of existing and extinct population samples clearly distinguished two genetically different groups, a northern and a southern clade. Because of the range contraction, the southern group is currently represented by very few extant populations. This southern edge range shift is thus causing the loss of a distinct component of the species genetic background.ConclusionsWe reveal a climate-correlated diversity loss below the species level, a process that could render the species more vulnerable to future environmental changes and affect its evolutionary potential. This is a remarkable case of genetic uniqueness of a vanishing cryptic genetic clade (southern clade).

Summary 1.Theory predicts that population structure and dynamics affect a population's capacity for adaptation to environmental change. For isolated, small and fragmented populations at the trailing edge of species distributions, loss of genetic diversity through random genetic drift may reduce adaptive potential and fitness levels for complex traits. This has important consequences for understanding population responses to, for example changing climate, but has rarely been tested in natural populations. 2. We measured the intertidal thermal environment and tidal exposure (emersion) times for natural populations of the intertidal seaweed Fucus serratus at the centre (southwest UK) and southern edge (northern Portugal) of its range in the Eastern Atlantic, and for a congener, F. vesiculosus , whose range extends further south to Morocco. Fitness-related traits of individuals at each location were measured in common garden experiments: physiological resilience to desiccation and heat shock (PSII quantum yield), and the molecular phenotype of the heat shock response (quantitative PCR of heat shock protein gene transcripts). 3. The realized thermal environment experienced by F. serratus was similar at the centre and southern edge of its distribution because the maximum shore height (and emersion period) was reduced in southern populations. For F. vesiculosus , thermal maxima were higher and occurred more frequently in the south, although maximum vertical height (emersion time) remained similar to central populations. 4. Edge populations of F. serratus were less resilient to desiccation and heat shock than central populations, and expression of heat shock genes was higher at the same temperature, suggesting greater cellular stress. In contrast, there was no evidence for physiological divergence in heat shock response in F. vesiculosus , and little variation in gene expression. 5. Synthesis. We provide evidence that compared with range-centre populations upper intertidal limits of F. serratus at the southern edge are 'pruned back' by abiotic stressors. Rather than being locally adapted, these small populations are less resilient to abiotic stresses and experience greater cellular stress during heat shock. These results suggest that ongoing climate forcing factors may threaten small, fragmented rear edge populations because of inherently reduced fitness and lower adaptive capacity relative to larger central populations.

Abstract. The North Atlantic intertidal community provides a rich set of organismal and environmental material for the study of ecological genetics. Clearly defined environmental gradients exist at multiple spatial scales: there are broad latitudinal trends in temperature, meso-scale changes in salinity along estuaries, and smaller scale gradients in desiccation and temperature spanning the intertidal range. The geology and geography of the American and European coasts provide natural replication of these gradients, allowing for population genetic analyses of parallel adaptation to environmental stress and heterogeneity. Statistical methods have been developed that provide genomic neutrality tests of population differentiation and aid in the process of candidate gene identification. In this paper, we review studies of marine organisms that illustrate associations between an environmental gradient and specific genetic markers. Such highly differentiated markers become candidate genes for adaptation to the environmental factors in question, but the functional significance of genetic variants must be comprehensively evaluated. We present a set of predictions about locus-specific selection across latitudinal, estuarine, and intertidal gradients that are likely to exist in the North Atlantic. We further present new data and analyses that support and contradict these simple selection models. Some taxa show pronounced clinal variation at certain loci against a background of mild clinal variation at many loci. These cases illustrate the procedures necessary for distinguishing selection driven by internal genomic vs. external environmental factors. We suggest that the North Atlantic intertidal community provides a model system for identifying genes that matter in ecology due to the clarity of the environmental stresses and an extensive experimental literature on ecological function. While these organisms are typically poor genetic and genomic models, advances in comparative genomics have provided access to molecular tools that can now be applied to taxa with well-defined ecologies. As many of the organisms we discuss have tight physiological limits driven by climatic factors, this synthesis of molecular population genetics with marine ecology could provide a sensitive means of assessing evolutionary responses to climate change.