Gene regulatory response to hyposalinity in the brown seaweed Fucus vesiculosus.

BACKGROUND: Rockweeds are among the most important foundation species of temperate rocky littoral shores. In the Baltic Sea, the rockweed Fucus vesiculosus is distributed along a decreasing salinity gradient from the North Atlantic entrance to the low-salinity regions in the north-eastern margins, thus, demonstrating a remarkable tolerance to hyposalinity. The underlying mechanisms for this tolerance are still poorly understood. Here, we exposed F. vesiculosus from two range-margin populations to the hyposaline (2.5 PSU - practical salinity unit) conditions that are projected to occur in the region by the end of this century as a result of climate change. We used transcriptome analysis (RNA-seq) to determine the gene expression patterns associated with hyposalinity acclimation, and examined the variation in these patterns between the sampled populations. RESULTS: Hyposalinity induced different responses in the two populations: in one, only 26 genes were differentially expressed between salinity treatments, while the other population demonstrated up- or downregulation in 3072 genes. In the latter population, the projected future hyposalinity induced an acute response in terms of antioxidant production. Genes associated with membrane composition and structure were also heavily involved, with the upregulation of fatty acid and actin production, and the downregulation of ion channels and alginate pathways. Changes in gene expression patterns clearly indicated an inhibition of the photosynthetic machinery, with a consequent downregulation of carbohydrate production. Simultaneously, energy consumption increased, as revealed by the upregulation of genes associated with respiration and ATP synthesis. Overall, the genes that demonstrated the largest increase in expression were ribosomal proteins involved in translation pathways. The fixation rate of SNP:s was higher within genes responding to hyposalinity than elsewhere in the transcriptome. CONCLUSIONS: The high fixation rate in the genes coding for salinity acclimation mechanisms implies strong selection for them. The among-population differentiation that we observed in the transcriptomic response to hyposalinity stress suggests that populations of F. vesiculosus may differ in their tolerance to future desalination, possibly as a result of local adaptation to salinity conditions within the Baltic Sea. These results emphasise the importance of considering interspecific genetic variation when evaluating the consequences of environmental change.

Smooth or smothering? The self-cleaning potential and photosynthetic effects of oil spill on arctic macro-algae Fucus distichus.

Due to increased sea transport and offshore gas and oil exploration, the Arctic is facing an unprecedented risk of marine oil spills. Although beached oil spills can lead to acute and chronic impacts on intertidal ecosystems, the effects of oil spills on macro-algae in Arctic ecosystems is lacking. Here, we assessed the effect and response of the tidal macro-algae Fucus distichus to oiling, i.e. self-cleaning potential by seawater wash and photosynthetic activity. Oiling with four oil types (ANS, Grane, IFO30 and MGO) was simulated by exposing F. distichus tips to oil. Oil removal half-times ranged between 0.8 - 4.5 days, indicating that oiling of macro-algae with the tested oils was short-term. Further, Grane oil mostly inhibited photosynthetic activity whereas oil from ANS, IFO30 and MGO stimulated it. The photosynthetic activity of F. distichus continued to be affected (inhibited or stimulated), even after oil on the tip surface was washed off.

Sensitivities to global change drivers may correlate positively or negatively in a foundational marine macroalga.

Ecological impact of global change is generated by multiple synchronous or asynchronous drivers which interact with each other and with intraspecific variability of sensitivities. In three near-natural experiments, we explored response correlations of full-sibling germling families of the seaweed Fucus vesiculosus towards four global change drivers: elevated CO2 (ocean acidification, OA), ocean warming (OW), combined OA and warming (OAW), nutrient enrichment and hypoxic upwelling. Among families, performance responses to OA and OW as well as to OAW and nutrient enrichment correlated positively whereas performance responses to OAW and hypoxia anti-correlated. This indicates (i) that families robust to one of the three drivers (OA, OW, nutrients) will also not suffer from the two other shifts, and vice versa and (ii) families benefitting from OAW will more easily succumb to hypoxia. Our results may imply that selection under either OA, OW or eutrophication would enhance performance under the other two drivers but simultaneously render the population more susceptible to hypoxia. We conclude that intraspecific response correlations have a high potential to boost or hinder adaptation to multifactorial global change scenarios.

Tolerance of Fucus vesiculosus exposed to diesel water-accommodated fraction (WAF) and degradation of hydrocarbons by the associated bacteria.

The viability and physiological state of brown macroalgae Fucus vesiculosus and its associated epiphytic bacteria exposed to diesel water-accommodated fraction (WAF), as well as the capacity of this association to deplete petroleum hydrocarbons (HCs) were experimentally tested. After a 6-day exposure treatment, the algal-surface associated bacteria were identified as primarily hydrocarbon-oxidising bacteria (HOB), and the algal-HOB association was able to deplete petroleum hydrocarbons from the diesel WAF by 80%. The HOB density on the algal surface exposed to diesel WAF was 350% higher compared to the control (i.e. HOB density on the algal surface exposed to ambient seawater), which suggest that they actively proliferated in the presence of hydrocarbons and most likely consumed hydrocarbons as their primary organic substrate. Exposure to diesel WAF did not affect the metabolic activity of F. vesiculosus. Higher lipid peroxidation was observed in F. vesiculosus exposed to diesel WAF while catalase concentration decreased only during the first day of exposure. Results suggest F. vesiculosus is tolerant to oil pollution and the algal-HOB association can efficiently deplete petroleum hydrocarbons in oil-contaminated seas.

Integrating experimental and distribution data to predict future species patterns.

Predictive species distribution models are mostly based on statistical dependence between environmental and distributional data and therefore may fail to account for physiological limits and biological interactions that are fundamental when modelling species distributions under future climate conditions. Here, we developed a state-of-the-art method integrating biological theory with survey and experimental data in a way that allows us to explicitly model both physical tolerance limits of species and inherent natural variability in regional conditions and thereby improve the reliability of species distribution predictions under future climate conditions. By using a macroalga-herbivore association (Fucus vesiculosus - Idotea balthica) as a case study, we illustrated how salinity reduction and temperature increase under future climate conditions may significantly reduce the occurrence and biomass of these important coastal species. Moreover, we showed that the reduction of herbivore occurrence is linked to reduction of their host macroalgae. Spatial predictive modelling and experimental biology have been traditionally seen as separate fields but stronger interlinkages between these disciplines can improve species distribution projections under climate change. Experiments enable qualitative prior knowledge to be defined and identify cause-effect relationships, and thereby better foresee alterations in ecosystem structure and functioning under future climate conditions that are not necessarily seen in projections based on non-causal statistical relationships alone.

Tolerance to climate change of the clonally reproducing endemic Baltic seaweed, Fucus radicans: is phenotypic plasticity enough?

To predict the effects of climate change, we first need information on both the current tolerance ranges of species and their future adaptive potential. Adaptive responses may originate either in genetic variation or in phenotypic plasticity, but the relative importance of these factors is poorly understood. Here, we tested the tolerance of Fucus radicans to the combination of hyposalinity and warming projected by climate models for 2070-2099. We measured the growth and survival responses of thalli in both current and future conditions, focusing on variations in tolerance among and within different clonal lineages. Survival was 32% lower in future than in current conditions, but the weight and length of the thalli which survived was respectively 267% and 178% higher when exposed to future conditions. The relatively high tolerance to the future conditions suggests that F. radicans is likely to persist in its current distributional range, which is limited to the Gulf of Bothia and Estonian coast in the Baltic Sea. Furthermore, this species may be able to expand its distribution southward and replace its congener F. vesiculosus, which, in previous studies, has not tolerated the future conditions as well. In addition, we discovered variation in tolerance to future conditions within one of the clonal lineages, which have been hitherto presumed to lack adaptive variation. The discovery of intra-clonal phenotypic plasticity means that this alga has the potential for adaptive responses to climate change, which may be the key to the future persistence of F. radicans in the Baltic Sea.

Living apart-together: Microhabitat differentiation of cryptic nematode species in a saltmarsh habitat.

Coexistence of highly similar species is at odds with ecological theory of competition; coexistence, then, requires stabilizing mechanisms such as differences in ecological niche. In the bacterivore nematode Litoditis marina species complex, which occurs associated with macro-algae, four cryptic lineages (Pm I-IV) co-occur in the field along the south-western coast and estuaries of The Netherlands. Here we investigate the temporal and/or spatial niche differentiation in their natural environment using a qPCR-based detection and relative quantification method. We collected different algal species (i.e. two Fucus species and Ulva sp.) and separated algal structures (i.e. receptacula, thalli, non-fertile tips and bladders) at different sampling months and times (i.e. twice per sampling month), to examine differences in microhabitat use between coexisting L. marina species. Results demonstrate that the cryptic species composition varied among different algal species and algal structures, which was also subject to temporal shifts. Pm I dominated on Fucus spp., Pm II showed dominance on Ulva sp., while Pm III overall had the lowest frequencies. Microhabitat partitioning was most pronounced between the two cryptic species which had similar microbiomes (Pm I and Pm II), and less so between the two species which had significantly different microbiomes (Pm I and Pm III), suggesting that species which share the same microhabitats may avoid competition through resource partitioning. The interplay of microhabitat differentiation and temporal dynamics among the cryptic species of L. marina implies that there is a complex interaction between biotic components and abiotic factors which contributes to their coexistence in the field.

Physiological responses to variations in grazing and light conditions in native and invasive fucoids.

Poor physiological acclimatization to climate change has led to shifts in the distributional ranges of various species and to biodiversity loss. However, evidence also suggests the relevance of non-climatic physical factors, such as light, and biotic factors, which may act in interactive or additive way. We used a mechanistic approach to evaluate the ecophysiological responses of four seaweed species (three dominant intertidal fucoids, Fucus serratus, Ascophyllum nodosum, Bifurcaria bifurcata, and the invasive Sargassum muticum) to different conditions of grazing, light irradiance and ultraviolet (UV) radiation. We performed a large-scale mesocosm experiment with a total of 800 individual thalli of macroalgae. The factorial experimental design included major algal traits, photoacclimation, nutrient stoichiometry and chemical defence as response variables. Few significant effects of the factors acting alone or in combination were observed, suggesting a good capacity for acclimatization in all four species. The significant effects were generally additive and there were no potentially deleterious synergistic effects between factors. Fucus serratus, a species currently undergoing a drastic contraction of its southern distribution limit in Europe, was the most strongly affected species, showing overall lower photosynthetic efficiency than the other species. The growth rate of F. serratus decreased when UV radiation was filtered out, but only in the presence of grazers. Moreover, more individuals of this species tended to reach maturity in the absence of grazers, and the nitrogen content of tissues decreased under full-spectrum light. Only the phlorotannin content of tissues of B. bifurcata and of exudates of A. nodosum, both slow-growing species, were positively affected by respectively removal of UVB radiation and the presence of grazers. The findings for S. muticum, a well-established invasive seaweed across European coasts, suggested similar physiological response of this fast-growing species to different levels of grazing activity and light quality/intensity. As expected, this species grew faster than the other species. Bifurcaria bifurcata and A. nodosum only showed minor effects of light quality and grazing on phlorotannins content, which suggests good resistance of these two long-lived species to the experimental conditions. Mechanistic approaches that are designed to analyse interactive effects of physical and biotic factors provide an understanding of physiological responses of species and help to improve the confidence of predictive distribution models.

Environmental and phenotypic heterogeneity of populations at the trailing range-edge of the habitat-forming macroalga Fucus serratus.

Empirical work on the dynamics of range limits of species distributions often lack replications of edge populations. We compared the local environment and performance of two groups of geographically peripheral populations of the foundation intertidal alga Fucus serratus L. at its southern range boundary in the NW Iberian Peninsula. Two populations were located on the Western Galician coast in large embayments or rias, and the other two on a Northern open coastal stretch in Lugo province. Sharp differences were detected in the local environment and performance of the two groups of populations. While recruitment was quite consistent throughout the year in rias, it was very limited and variable in Lugo. Furthermore, thalli from rias were severely damaged following their transplantation in Lugo, and poor conditions of local adult plants were detected there in subsequent years. These results suggest a higher vulnerability of Lugo populations under new climate conditions, while western rias, strongly influenced by upwelling events, feasibly act as contemporary refugia for this species. If sustained over time, these refugia may mitigate the retreat of the seaweed' rear-edge predicted by large spatial scale models.

Tolerance and potential for adaptation of a Baltic Sea rockweed under predicted climate change conditions.

Climate change is threating species' persistence worldwide. To predict species responses to climate change we need information not just on their environmental tolerance but also on its adaptive potential. We tested how the foundation species of rocky littoral habitats, Fucus vesiculosus, responds to combined hyposalinity and warming projected to the Baltic Sea by 2070-2099. We quantified responses of replicated populations originating from the entrance, central, and marginal Baltic regions. Using replicated individuals, we tested for the presence of within-population tolerance variation. Future conditions hampered growth and survival of the central and marginal populations whereas the entrance populations fared well. Further, both the among- and within-population variation in responses to climate change indicated existence of genetic variation in tolerance. Such standing genetic variation provides the raw material necessary for adaptation to a changing environment, which may eventually ensure the persistence of the species in the inner Baltic Sea.

The role of chemical antifouling defence in the invasion success of Sargassum muticum: A comparison of native and invasive brown algae.

Competition and fouling defence are important traits that may facilitate invasions by non-indigenous species. The 'novel weapons hypothesis' (NWH) predicts that the invasive success of exotic species is closely linked to the possession of chemical defence compounds that the recipient community in the new range is not adapted to. In order to assess whether chemical defence traits contribute to invasion success, anti-bacterial, anti-quorum sensing, anti-diatom, anti-larval and anti-algal properties were investigated for the following algae: a) the invasive brown alga Sargassum muticum from both, its native (Japan) and invasive (Germany) range, b) the two non- or weak invasive species Sargassum fusiforme and Sargassum horneri from Japan, and c) Fucus vesiculosus, a native brown alga from Germany. Crude and surface extracts and lipid fractions of active extracts were tested against common fouling organisms and zygotes of a dominant competing brown alga. Extracts of the native brown alga F. vesiculosus inhibited more bacterial strains (75%) than any of the Sargassum spp. (17 to 29%). However, Sargassum spp. from Japan exhibited the strongest settlement inhibition against the diatom Cylindrotheca closterium, larvae of the bryozoan Bugula neritina and zygotes of the brown alga F. vesiculosus. Overall, extracts of S. muticum from the invasive range were less active compared to those of the native range suggesting an adaptation to lower fouling pressure and competition in the new range resulting in a shift of resource allocation from costly chemical defence to reproduction and growth. Non-invasive Sargassum spp. from Japan was equally defended against fouling and competitors like S. muticum from Japan indicating a necessity to include these species in European monitoring programs. The variable antifouling activity of surface and crude extracts highlights the importance to use both for an initial screening for antifouling activity.

Light Sheet Microscopy Imaging of Light Absorption and Photosynthesis Distribution in Plant Tissue.

In vivo variable chlorophyll fluorescence measurements of photosystem II (PSII) quantum yields in optically dense systems are complicated by steep tissue light gradients due to scattering and absorption. Consequently, externally measured effective PSII quantum yields may be composed of signals derived from cells differentially exposed to actinic light, where cells located deeper inside tissues receive lower irradiance than cells closer to the surface and can display distinct photophysiological status. We demonstrate how measured distributions of PSII quantum yields in plant tissue change under natural tissue light gradients as compared with conventionally measured quantum yields with even exposure to actinic light. This was achieved by applying actinic irradiance perpendicular to one side of thallus cross sections of the aquatic macrophyte Fucus vesiculosus with laser light sheets of defined spectral composition, while imaging variable chlorophyll fluorescence from cross sections with a microscope-mounted pulse amplitude-modulated imaging system. We show that quantum yields are highly affected by light gradients and that traditional surface-based variable chlorophyll fluorescence measurements result in substantial underestimations and/or overestimations, depending on incident actinic irradiance. We present a method for using chlorophyll fluorescence profiles in combination with integrating sphere measurements of reflectance and transmittance to calculate depth-resolved photon absorption profiles, which can be used to correct apparent PSII electron transport rates to photons absorbed by PSII. Absorption profiles of the investigated aquatic macrophyte were different in shape from what is typically observed in terrestrial leaves, and based on this finding, we discuss strategies for optimizing photon absorption via modulation of the structural organization of phytoelements according to in situ light environments.

Translocation of isotopically distinct macroalgae: A route to low-cost biomonitoring?

Nitrogen stable isotope ratios (δ15N) in macroalgae are often used to identify sources of nitrogenous pollution in fluvial and estuarine settings. This approach assumes that the macroalgal δ15N is representative of the sources of the pollution averaged over a timespan in the order of days to weeks. The preferential uptake of a particular nitrogen compound or potential for fractionation in the water column or during uptake and assimilation by the macroalgae could make this assumption invalid. Laboratory studies were therefore performed to investigate the uptake and assimilation of both nitrate and ammonium at a variety of concentrations using the vegetative (non-fertile) tips of the brown macroalgae, Fucus vesiculosus. Nitrate appeared to fractionate at high concentrations, and was found to be taken up more rapidly than ammonia; within 13 days, the macroalgae tips were in isotopic equilibrium with the nitrate solution at 500 µM. These experiments were complemented by an investigation involving the translocation of macroalgae collected from a site enriched in 15N relative to natural levels (Staithes, UK), to the River Tees, Middlesbrough (UK), a site depleted in 15N relative to natural levels. The nitrogen isotope signature shifted by ∼50% within 7 days, with samples deployed nearer the surface subject to greater change. These findings suggest that the translocation of macroalgae with isotopically distinct signatures can be used as a rapid, cost-efficient method for nitrogen biomonitoring in estuarine environments.

Reciprocal transplants support a plasticity-first scenario during colonisation of a large hyposaline basin by a marine macro alga.

BACKGROUND: Establishing populations in ecologically marginal habitats may require substantial phenotypic changes that come about through phenotypic plasticity, local adaptation, or both. West-Eberhard's "plasticity-first" model suggests that plasticity allows for rapid colonisation of a new environment, followed by directional selection that develops local adaptation. Two predictions from this model are that (i) individuals of the original population have high enough plasticity to survive and reproduce in the marginal environment, and (ii) individuals of the marginal population show evidence of local adaptation. Individuals of the macroalga Fucus vesiculosus from the North Sea colonised the hyposaline (≥2-3‰) Baltic Sea less than 8000 years ago. The colonisation involved a switch from fully sexual to facultative asexual recruitment with release of adventitious branches that grow rhizoids and attach to the substratum. To test the predictions from the plasticity-first model we reciprocally transplanted F. vesiculosus from the original population (ambient salinity 24‰) and from the marginal population inside the Baltic Sea (ambient salinity 4‰). We also transplanted individuals of the Baltic endemic sister species F. radicans from 4 to 24‰. We assessed the degree of plasticity and local adaptation in growth and reproductive traits after 6 months by comparing the performance of individuals in 4 and 24‰. RESULTS: Branches of all individuals survived the 6 months period in both salinities, but grew better in their native salinity. Baltic Sea individuals more frequently developed asexual traits while North Sea individuals initiated formation of receptacles for sexual reproduction. CONCLUSIONS: Marine individuals of F. vesiculosus are highly plastic with respect to salinity and North Sea populations can survive the extreme hyposaline conditions of the Baltic Sea without selective mortality. Plasticity alone would thus allow for an initial establishment of this species inside the postglacial Baltic Sea at salinities where reproduction remains functional. Since establishment, the Baltic Sea populations have evolved adaptations to extreme hyposaline waters and have in addition evolved asexual recruitment that, however, tends to impede local adaptation. Overall, our results support the "plasticity-first" model for the initial colonisation of the Baltic Sea by Fucus vesiculosus.

Diffusion or advection? Mass transfer and complex boundary layer landscapes of the brown alga Fucus vesiculosus.

The role of hyaline hairs on the thallus of brown algae in the genus Fucus is long debated and several functions have been proposed. We used a novel motorized set-up for two-dimensional and three-dimensional mapping with O2 microsensors to investigate the spatial heterogeneity of the diffusive boundary layer (DBL) and O2 flux around single and multiple tufts of hyaline hairs on the thallus of Fucus vesiculosus. Flow was a major determinant of DBL thickness, where higher flow decreased DBL thickness and increased O2 flux between the algal thallus and the surrounding seawater. However, the topography of the DBL varied and did not directly follow the contour of the underlying thallus. Areas around single tufts of hyaline hairs exhibited a more complex mass-transfer boundary layer, showing both increased and decreased thickness when compared with areas over smooth thallus surfaces. Over thallus areas with several hyaline hair tufts, the overall effect was an apparent increase in the boundary layer thickness. We also found indications for advective O2 transport driven by pressure gradients or vortex shedding downstream from dense tufts of hyaline hairs that could alleviate local mass-transfer resistances. Mass-transfer dynamics around hyaline hair tufts are thus more complex than hitherto assumed and may have important implications for algal physiology and plant-microbe interactions.

Chemotactic movement in sperm of the oogamous brown algae, Saccharina japonica and Fucus distichus.

In oogamous species of brown algae such as Saccharina japonica and Fucus distichus, the sperm possess an unusual long posterior flagellum, which oscillates actively and produces a propulsive force during swimming. In this study, we quantitatively analyzed the effect of chemotactic responses on sperm swimming and flagellar waveforms by high-speed video recordings. We found that the thigmotactic response to the chemo-attractant was not enhanced during chemotactic swimming and that the swimming velocity of sperm did not decrease. As concentration of the chemo-attractant decreased, the sperm performed drastic U-turn movements, which was caused by a rapid and large bend of the posterior flagellum. Unilateral bending of the posterior flagellum when sensing a decrease in the concentration of the chemo-attractant may be a common response in male gametes during fertilization of brown algae both oogamous and isogamous species.

Future warming and acidification effects on anti-fouling and anti-herbivory traits of the brown alga Fucus vesiculosus (Phaeophyceae).

Human-induced ocean warming and acidification have received increasing attention over the past decade and are considered to have substantial consequences for a broad range of marine species and their interactions. Understanding how these interactions shift in response to climate change is particularly important with regard to foundation species, such as the brown alga Fucus vesiculosus. This macroalga represents the dominant habitat former on coastal rocky substrata of the Baltic Sea, fulfilling functions essential for the entire benthic community. Its ability to withstand extensive fouling and herbivory regulates the associated community and ecosystem dynamics. This study tested the interactive effects of future warming, acidification, and seasonality on the interactions of a marine macroalga with potential foulers and consumers. F. vesiculosus rockweeds were exposed to different combinations of conditions predicted regionally for the year 2100 (+∆5°C, +∆700 µatm CO2 ) using multifactorial long-term experiments in novel outdoor benthic mesocosms ("Benthocosms") over 9-12-week periods in four seasons. Possible shifts in the macroalgal susceptibility to fouling and consumption were tested using consecutive bioassays. Algal susceptibility to fouling and grazing varied substantially among seasons and between treatments. In all seasons, warming predominantly affected anti-fouling and anti-herbivory interactions while acidification had a subtle nonsignificant influence. Interestingly, anti-microfouling activity was highest during winter under warming, while anti-macrofouling and anti-herbivory activities were highest in the summer under warming. These contrasting findings indicate that seasonal changes in anti-fouling and anti-herbivory traits may interact with ocean warming in altering F. vesiculosus community composition in the future.

Nitrate uptake varies with tide height and nutrient availability in the intertidal seaweed Fucus vesiculosus.

Intertidal seaweeds must cope with a suite of stressors imposed by aerial exposure at low tide, including nutrient limitation due to emersion. Seaweeds can access nutrients only when submerged, so individuals living higher compared to lower on the shore may have adaptations allowing them to acquire sufficient amounts of nutrients to survive and maintain growth. Using a combination of observations and experiments, we aimed to identify intraspecific variation in nitrate uptake rates across the intertidal distribution of F. vesiculosus, as well as test for acclimation in response to a change in tide height. We replicated our study at sites spanning nearly the entire Gulf of Maine coastline, to examine how local environmental variability may alter intraspecific variation in nitrate uptake. We found that average nitrate uptake rates were ~18% higher in upper compared to lower intertidal Fucus vesiculosus. Furthermore, we found evidence for both acclimation and adaptation to tide height during a transplant experiment. F. vesiculosus transplanted from the lower to the upper intertidal zone was characterized by increased nitrate uptake, but individuals transplanted from the upper to the lower intertidal zone retained high uptake rates. Our observations differed among Gulf of Maine regions and among time points of our study. Importantly, these differences may reflect associations between nitrate uptake rates and abiotic environmental conditions and seaweed nutrient status. Our study highlights the importance of long-term variation in ambient nutrient supply in driving intraspecific variation of seaweeds across the intertidal gradient and local and seasonal variation in ambient nutrient levels in mediating intraspecific differences.

Temporal windows of reproductive opportunity reinforce species barriers in a marine broadcast spawning assemblage.

Prezygotic isolating mechanisms act to limit hybridization and maintain the genetic identity of closely-related species. While synchronous intraspecific spawning is a common phenomenon amongst marine organisms and plays an important role in reproductive success, asynchronous spawning between potentially hybridizing lineages may also be important in maintaining species boundaries. We tested this hypothesis by comparing reproductive synchrony over daily to hourly timescales in a sympatric assemblage of intertidal fucoid algae containing selfing hermaphroditic (Fucus spiralis and Fucus guiryi) and dioecious (Fucus vesiculosus and Fucus serratus) species. Our results confirm that gametes are released on semi-lunar cycles in all species. However, sister species with different mating systems showed asynchronous spawning at finer circadian timescales, thus providing evidence for a partial reproductive barrier between hermaphroditic and dioecious species. Finally, our data also emphasize the ecological, developmental, and/or physiological constraints that operate to restrict reproduction to narrow temporal windows of opportunity in the intertidal zone and more generally the role of ecological factors in marine speciation.

Home advantage? Decomposition across the freshwater-estuarine transition zone varies with litter origin and local salinity.

Expected increases in the frequency and intensity of storm surges and river flooding may greatly affect the relative salinity of estuarine environments over the coming decades. In this experiment we used detritus from three contrasting environments (marine Fucus vesiculosus; estuarine Spartina anglica; terrestrial Quercus robur) to test the prediction that the decomposition of the different types of litter would be highest in the environment with which they are associated. Patterns of decomposition broadly fitted our prediction: Quercus detritus decomposed more rapidly in freshwater compared with saline conditions while Fucus showed the opposite trend; Spartina showed an intermediate response. Variation in macro-invertebrate assemblages was detected along the salinity gradient but with different patterns between estuaries, suggesting that breakdown rates may be linked in part to local invertebrate assemblages. Nonetheless, our results suggest that perturbation of salinity gradients through climate change could affect the process of litter decomposition and thus alter nutrient cycling in estuarine transition zones. Understanding the vulnerability of estuaries to changes in local abiotic conditions is important given the need to better integrate coastal proceses into a wider management framework at a time when coastlines are increasingly threatened by human activities.