Kelp forests are declining in many regions globally with climatic perturbations causing shifts to alternate communities and significant ecological and economic loss. Range edge populations are often at most risk and are often only sustained through localised areas of upwelling or on deeper reefs. Here we document the loss of kelp forests ( Ecklonia radiata ) from the Sultanate of Oman, the only confirmed northern hemisphere population of this species. Contemporary surveys failed to find any kelp in its only known historical northern hemisphere location, Sadah on the Dhofar coast. Genetic analyses of historical herbarium specimens from Oman confirmed the species to be E. radiata and revealed the lost population contained a common CO1 haplotype found across South Africa, Australia and New Zealand suggesting it once established through rapid colonisation throughout its range. However, the Omani population also contained a haplotype that is found nowhere else in the extant southern hemisphere distribution of E. radiata . The loss of the Oman population could be due to significant increases in the Arabian Sea temperature over the past 40 years punctuated by suppression of coastal upwelling. Climate-mediated warming is threatening the persistence of temperate species and precipitating loss of unique genetic diversity at lower latitudes.

Understanding the factors and processes that shape intra-specific sensitivity to heat stress is fundamental to better predicting the vulnerability of benthic species to climate change. Here, we investigate the response of a habitat-forming Mediterranean octocoral, the red gorgonian Paramuricea clavata (Risso, 1826) to thermal stress at multiple biological and geographical scales. Samples from eleven P. clavata populations inhabiting four localities separated by hundreds to more than 1500 km of coast and with contrasting thermal histories were exposed to a critical temperature threshold (25 °C) in a common garden experiment in aquaria. Ten of the 11 populations lacked thermotolerance to the experimental conditions provided (25 days at 25 °C), with 100% or almost 100% colony mortality by the end of the experiment. Furthermore, we found no significant association between local average thermal regimes nor recent thermal history (i.e., local water temperatures in the 3 months prior to the experiment) and population thermotolerance. Overall, our results suggest that local adaptation and/or acclimation to warmer conditions have a limited role in the response of P. clavata to thermal stress. The study also confirms the sensitivity of this species to warm temperatures across its distributional range and questions its adaptive capacity under ocean warming conditions. However, important inter-individual variation in thermotolerance was found within populations, particularly those exposed to the most severe prior marine heatwaves. These observations suggest that P. clavata could harbor adaptive potential to future warming acting on standing genetic variation (i.e., divergent selection) and/or environmentally-induced phenotypic variation (i.e., intra- and/or intergenerational plasticity).

The copepod Calanus glacialis plays a key role in the Arctic pelagic ecosystem. Despite its ecological importance and ongoing climate changes, limited knowledge at the genomic level has hindered the understanding of the molecular processes underlying environmental stress responses and ecological adaptation. Transcriptome data was generated from an experiment with C. glacialis copepodite (CV) subjected to five different temperatures. We obtained a total of 512,352 high-quality 454 pyrosequencing reads, which were assembled into 55,562 contigs distributed in 128 KEGG pathways. Functional analysis revealed numerous genes related to diverse biological functions and processes, including members of all major conserved signaling pathways. Comparative analysis of acclimated individuals to experimental temperatures has provided information about gene variations observed in several pathways (e.g. genes involved in energy, lipid and amino acid metabolism were shown to be down-regulated with increasing temperatures). These mRNA sequence resources will facilitate further studies on genomics and physiology-driven molecular processes in C. glacialis and related species.

A fundamental goal in ecology is to understand distribution and abundance of species. Peripheral populations inhabiting the trailing‐edge of a species' distribution may carry considerable ecological and evolutionary value yet being most threatened under predicted climate change scenarios. However, the nature of species distributional limits and the ecological and genetic implications of living at low latitude rear edges remain unclear. The assumption that population abundance declines towards range edges, where environmental selective pressure increases, is the basis of a wide range of ecological and evolutionary predictions. Empirical data have provided contrasting evidence about the consequences of living at low latitude distributional limits, raising concerns regarding their generality and highlighting the need for combined multidisciplinary tests. Here, we provide a long‐term, comprehensive evaluation of latitudinal patterns in genetic diversity, demographic, morphological and life history traits in the marine macroalga Fucus guiryi.The center to edge transition was mirrored by increasing water and air temperature gradients, with sea surface temperature of coldest months being the most relevant influence on F. guiryi traits. Overall, we identified a strong correlation between ecological data, collected over one year at bimonthly intervals, and distance to the range center. We found decreasing population and individual size towards distributional margins. Similarly, reproductive capacity, threshold size for reproduction, density of reproductive population and recruitment rates showed a core to edge reduction. Temporal variability of individual reproductive effort and recruitment rates did not conform to the general pattern. In contrast, population genetic data did not show a core‐edge gradient, as gene diversity and allelic richness were not significantly lower at edge populations, contradicting predictions of higher drift and bottlenecks for smaller edge populations.The contrasting support provided by genetic and ecological data highlights the need to combine multiple and cross‐disciplinary evidence for a comprehensive understanding of ecological and evolutionary mechanisms linked to species ranges.

The climate-driven dynamics of species ranges is a critical research question in evolutionary ecology. We ask whether present intraspecific diversity is determined by the imprint of past climate. This is an ongoing debate requiring interdisciplinary examination of population genetic pools and persistence patterns across global ranges. Previously, contrasting inferences and predictions have resulted from distinct genomic coverage and/or geographical information. We aim to describe and explain the causes of geographical contrasts in genetic diversity and their consequences for the future baseline of the global genetic pool, by comparing present geographical distribution of genetic diversity and differentiation with predictive species distribution modelling (SDM) during past extremes, present time and future climate scenarios for a brown alga, Fucus vesiculosus. SDM showed that both atmospheric and oceanic variables shape the global distribution of intertidal species, revealing regions of persistence, extinction and expansion during glacial and postglacial periods. These explained the distribution and structure of present genetic diversity, consisting of differentiated genetic pools with maximal diversity in areas of long-term persistence. Most of the present species range comprises postglacial expansion zones and, in contrast to highly dispersive marine organisms, expansions involved only local fronts, leaving distinct genetic pools at rear edges. Besides unravelling a complex phylogeographical history and showing congruence between genetic diversity and persistent distribution zones, supporting the hypothesis of niche conservatism, range shifts and loss of unique genetic diversity at the rear edge were predicted for future climate scenarios, impoverishing the global gene pool.

Understanding the extent to which neutral processes and adaptive divergence shape the spatial structure of natural populations is a major goal in evolutionary biology and is especially important for the identification of significant levels of biodiversity. Our results identified replicated habitat-specific (adaptive) phenotypic divergence in the brown macroalga Fucus vesiculosus that is independent of population (neutral) genetic structure. F. vesiculosus inhabits contiguous and contrasting marine to estuarine intertidal habitats. Combining analyses of genetic and phenotypic traits of populations living under differential selective regimes (estuaries and open coast), we investigated levels of neutral genetic differentiation and adaptive physiological responses to emersion stress. In southwest England (SW UK) and northern Iberia (N. Iberia), populations living in estuaries and marine coastal habitats were genetically characterized at six microsatellite loci. In N. Iberia, two clades with limited admixture were recovered, each including one open coast site and the adjacent estuarine location. In contrast, SW UK samples clustered according to habitat and formed three distinct groups of genotypes; one including the two open coast locations and the other two representing each of the estuarine sites. Temperature loggers revealed distinct emersion regimes that characterized each habitat type independently of the region, while water and air temperature profiles showed site-specific trends. Despite acclimation under usual conditions, trait means of emersion stress resilience showed a strong phenotypic divergence between habitats, consistent with environmental dines in exposure time observed in the different habitats. We demonstrate that neutral genetic clusters do not reflect locally adapted population units. Our results identified replicated habitat-specific (adaptive) phenotypic divergence that is independent of population (neutral) genetic structure in E vesiculosus. The significance of such findings extends beyond the theoretical evolutionary and ecological interest of discovering parallel adaptive responses to the broader implications for conservation of intraspecific biodiversity. (c) 2013 Elsevier Ltd. All rights reserved.

The effect of temperature on marine phytoplankton growth strategies, metabolism and composition is studied using a range of techniques. This approach indicates that temperature plays a previously unrecognized, critical role in resource allocation and marine phytoplankton elemental ratios, with implications for biogeochemical cycling.

We examined 733 individuals of Fucusspiralis from 21 locations and 1093 Fucusvesiculosus individuals from 37 locations throughout their northern hemisphere ranges using nuclear and mitochondrial markers. Three genetic entities of F. spiralis were recovered. In northern and sympatric populations, the presence of "F. spiralis Low" in the mid-intertidal and "F. spiralis High" in the high-intertidal was confirmed and both co-occurred with the sister species F. vesiculosus. The third and newly-discovered entity, "F. spiralis South", was present mainly in the southern range, where it did not co-occur with F. vesiculosus. The South entity diverged early in allopatry, then hybridized with F. vesiculosus in sympatry to produce F. spiralis Low. Ongoing parallel evolution of F. spiralis Low and F. spiralis High is most likely due to habitat preference/local selection and maintained by preferentially selfing reproductive strategies. Contemporary populations of F. spiralis throughout the North Atlantic stem from a glacial refugium around Brittany involving F. spiralis High; F. spiralis South was probably unaffected by glacial episodes. Exponential population expansion for F. vesiculosus began during the Cromer and/Holstein interglacial period (300,000-200,000 yrs BP). Following the last glacial maximum (30,000-22,000 yrs BP), a single mtDNA haplotype from a glacial refugium in SW Ireland colonized Scandinavia, the Central Atlantic islands, and the W Atlantic.

The timing of gamete release by fucoid algae, although known to be restricted to calm days is not clearly understood within a circadian time scale. The need for externally fertilizing species to avoid gamete dilution suggests that in wave-exposed areas spawning may occur during particular tidal phases. However, this may differ between mating systems, as selfing species may be less affected by gamete dilution. In this study, two different approaches were used to determine when egg release occurs during the tidal cycle in two sister species with different mating systems. First, egg settlement of Fucus vesiculosus (dioecious) and Fucus spiralis (selfing hermaphrodite) was quantified on removable substrates (egg settlement disks) every day for 2 months and settlement patterns were used to statistically estimate the radius of a circle that would encompass 99% of each patch of settled eggs (the egg dispersal radius). Also, egg release was quantified every 2 h during the tidal cycle. A significantly larger egg dispersal radius (P < 0.02) was found for F. spiralis than F. vesiculosus, and this difference was somewhat site dependent with a greater difference between species in exposed sites. The egg dispersal radius was negatively correlated with significant wave height and positively correlated with sea surface temperature for both the species (P < 0.05), with a greater effect of both the factors for F. spiralis than for F. vesiculosus. Egg release during the tidal cycle was variable between species and experiments, with F. vesiculosus releasing more eggs, later in the day, and at a lower tide, than F. spiralis, which released fewer eggs, throughout the day and at all tides. The dioecious species, F. vesiculosus, may have developed a specific adaptation for timing the egg release to periods when emersed in exposed habitats to avoid rapid dilution of gametes that require outcrossing for fertilization. On the other hand, egg release for F. spiralis, which can self-fertilize, occurred both when emersed and immersed, suggesting this species has developed less synchrony with specific environmental factors. Site dependence also suggests local wave conditions can modulate timing of release. The data are consistent with the relaxation of the selective constraints of water motion on fertilization success in a selfing hermaphrodite, relative to an obligate outcrossing species. Results support the idea that species with different mating systems evolve different sensitivities to environmental cues for gamete release with specific implications for inbreeding and successful external fertilization in the ocean.

The dwarf seagrass Zostera noltii is an important primary producer in Atlantic coastal ecosystems from Mauritania to southern Norway and the Mediterranean Sea. Sessile intertidal organisms existing at the interface between marine and terrestrial environments may be particularly vulnerable to environmental change. In this study, we asked how near to thermal tolerance limits natural populations of Z. noltii are in the Ria Formosa coastal lagoon system in southern Portugal. We recorded the maximum temperatures in the Ria Formosa during the 2007 summer, and conducted experiments to determine the sub-lethal temperature of Z. noltii shoots sampled at two sites located at different tidal heights. Mortality rates and photosynthetic performance were recorded within a range of heat shock temperatures between 35 and 41°C. Survival was recorded ≤37°C, while higher temperatures led to a sudden drop in photosynthetic capacity followed by mortality (shoot loss) that occurred more rapidly with increasing temperatures. At 39°C and above, the rate of shoot mortality in both sites was close to 100%, occurring between 5 and 13 days after the heat shock. Survival was ca. 95 and 90% at 35 and 37°C, respectively. From these results for Z. noltii populations in the Ria Formosa we estimated sub-lethal temperature to be approximately 38°C for Z. noltii, close to the maximum of 36°C recorded in the summer 2007. Considering predicted trajectories in the coming decades, these results raise concern as to the future viability of intertidal Z. noltii populations near the southernmost edge of their distribution.

Differential thermal analysis (DTA) was used to measure the freezing temperature of nine species of red brown intertidal macroalgae from the coast of Maine, USA in 1991. Using slow and rapid cooling rates approximating those found in the field for Ascophyllum nodosum (L.) Le Jol. we found that, for a given rate, the freezing points of all species were similar: -7.06 to -8.02°C for slow cooling (ca. 0.25°C min-1) and -3.42 to -4.56°C for rapid cooling (ca. 5.0°C min-1). In the low shore species, Fucus evanescens C. Ag., photosynthesis was inhibited to a greater extent when plants were frozen or thawed rapidly than after slow freezing or thawing. However, in the upper shore species, F. spiralis (L.), photosynthesis recovered rapidly and completely regardless of freezing rate. Rapidly frozen F. evanescens also experienced greater loss of plasmalemmal integrity, evidenced by a greater loss of cellular contents on re-immersion, than those frozen slowly. Light-limited photosynthesis following freezing was more severely inhibited than light-saturated photosynthesis. Respiration was generally enhanced immediately after freezing, but then declined to rates below those of unfrozen controls within 2 h following re-immersion, with control rates of respiration being achieved after a 24 h recovery period. Our data suggest that the physiological consequences of winter emersion at sub-zero temperatures may vary widely between individual plants of freezing-susceptible species, due to the wide variations in freezing rate associated with microhabitat effects.