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The impact of colonization by Sargassum muticum (Phaeophyta: Fucales) on the resident macroalgal assemblage in tidepools was investigated in northern Spain. Sargassum muticum was allowed to colonize and spread in some tidepools, and was periodically removed in others. Changes in the abundance of macrophytes were recorded at three sites over two years. The colonization by S. muticum and the subsequent changes induced on the native assemblage was extremely variable among sites. Where the invader colonized profusely, it induced changes in the macroalgal composition of tidepools. The leathery group of macrophytes was significantly affected and changes were also observed in the foliose group. It was suggested that S. muticum may take advantage of their perennial and opportunistic traits in the interaction with groups of algae with different survival strategies. However, the spatial patchiness of the colonization may mitigate the impact of the invasion at local and regional scales.

Species distribution models (SDM) are a useful tool for predicting species range shifts in response to global warming. However, they do not explore the mechanisms underlying biological processes, making it difficult to predict shifts outside the environmental gradient where the model was trained. In this study, we combine correlative SDMs and knowledge on physiological limits to provide more robust predictions. The thermal thresholds obtained in growth and survival experiments were used as proxies of the fundamental niches of two foundational marine macrophytes. The geographic projections of these species' distributions obtained using these thresholds and existing SDMs were similar in areas where the species are either absent-rare or frequent and where their potential and realized niches match, reaching consensus predictions. The cold-temperate foundational seaweed Himanthalia elongata was predicted to become extinct at its southern limit in northern Spain in response to global warming, whereas the occupancy of southern-lusitanic Bifurcaria bifurcata was expected to increase. Combined approaches such as this one may also highlight geographic areas where models disagree potentially due to biotic factors. Physiological thresholds alone tended to over-predict species prevalence, as they cannot identify absences in climatic conditions within the species' range of physiological tolerance or at the optima. Although SDMs tended to have higher sensitivity than threshold models, they may include regressions that do not reflect causal mechanisms, constraining their predictive power. We present a simple example of how combining correlative and mechanistic knowledge provides a rapid way to gain insight into a species' niche resulting in consistent predictions and highlighting potential sources of uncertainty in forecasted responses to climate change.

Climate change is driving species range shifts worldwide. However, physiological responses related to distributional changes are not fully understood. Oceanographers have reported an increase in ocean temperature in the northwest Iberian Peninsula that is potentially related to the decline in some cold-temperate intertidal macroalgae in the Cantabrian Sea, namely Fucus serratus. Low tide stress could also play a role in this decline. We performed one mensurative (in situ) and two manipulative (in culture) experiments designed to evaluate the interactive effects of some physical factors. The first experiment analysed field response to low tide stress in marginal (mid-Cantabrian Sea and northern Portugal) versus central (Galicia) populations of F. serratus. Then a second experiment was performed that utilized either harsh or mild summer conditions of atmospheric temperature, irradiance, humidity, and wind velocity to compare the responses of individuals from one marginal and one central population to low tide stress. Finally, the combined effect of sea temperature and the other factors was evaluated to detect interactive effects. Changes in frond growth, maximal photosynthetic quantum yield (F(v)/F(m)), temperature, and desiccation were found. Three additive factors (solar irradiation, ocean and air temperatures) were found to drive F. serratus distribution, except under mildly humid conditions that ameliorated atmospheric thermal stress (two additive factors). Mid-Cantabrian Sea temperatures have recently increased, reaching the inhibitory levels suggested in this study of F. serratus. We also expect an additive secondary contribution of low tide stress to this species decline. On the northern Portugal coast, ocean warming plus low tide stress has not reached this species' inhibition threshold. No significant differential responses attributed to the population of origin were found. Mechanistic approaches that are designed to analyse the interactive effects of physical stressors may improve the levels of confidence in predicted range shifts of species.

Aim  Because intertidal organisms often live close to their physiological tolerance limits, they are potentially sensitive indicators of climate‐driven changes in the environment. The goals of this study were to assess the effect of climatic and non‐climatic factors on the geographical distribution of intertidal macroalgae, and to predict future distributions under different climate‐warming scenarios. Location  North‐western Iberian Peninsula, southern Europe. Methods  We developed distribution models for six ecologically important intertidal seaweed species. Occurrence and microhabitat data were sampled at 1‐km2 resolution and analysed with climate variables measured at larger spatial scales. We used generalized linear models and applied the deviance and Bayesian information criterion to model the relationship between environmental variables and the distribution of each target species. We also used hierarchical partitioning (HP) to identify predictor variables with higher independent explanatory power. Results  The distributions of Himanthalia elongata and Bifurcaria bifurcata were correlated with measures of terrestrial and marine climate, although in opposite directions. Model projections under two warming scenarios indicated the extinction of the former at a faster rate in the Cantabrian Sea (northern Spain) than in the Atlantic (west). In contrast, these models predicted an increase in the occurrence of B. bifurcata in both areas. The occurrences of Ascophyllum nodosum and Pelvetia canaliculata, species showing rather static historical distributions, were related to specific non‐climatic environmental conditions and locations, such as the location of sheltered sites. At the southernmost distributional limit, these habitats may present favourable microclimatic conditions or provide refuges from competitors or natural enemies. Model performances for Fucus vesiculosus and F. serratus were similar and poor, but several climatic variables influenced the occurrence of the latter in the HP analyses. Main conclusions  The correlation between species distributions and climate was evident for two species, whereas the distributions of the others were associated with non‐climatic predictors. We hypothesize that the distribution of F. serratus responds to diverse combinations of factors in different sections of the north‐west Iberian Peninsula. Our study shows how the response of species distributions to climatic and non‐climatic variables may be complex and vary geographically. Our analyses also highlight the difficulty of making predictions based solely on variation in climatic factors measured at coarse spatial scales.

Understanding the factors determining geographic ranges and range shifts of species is a central issue in ecology and evolutionary biology. Research addressing distributional borders from a demographic perspective frequently focused on reproductive traits, finding reproduction reductions or failure at the range margin. However, some of the observed changes in marginal locations could be the result of adaptive adjustments to local, unfavourable conditions, though they have been rarely interpreted from this point of view. In this study we investigated the reproductive patterns of the seaweed Fucus serratus in central and southern marginal locations (SW UK, N Spain) over a 3‐yr period. Our main goals were: 1) to determine the spatial (centre‐margin) and temporal variation in reproductive traits and 2) to test if this variation fits with life‐history predictions for stressful environments. Threshold size for reproduction declined at the range margin, in accordance with life‐history predictions. Nevertheless, we also observed parallel drastic reductions in the percentage of reproductives, reproductive allocation and plant size. The reproductive capacity of marginal locations was thus dramatically reduced in relation to central ones. Furthermore, the decline became more pronounced over the study period. Our results suggests that the viability of marginal populations is at risk. This situation clearly differs from the pattern observed during the last decade. At that time, the species was able to growth and reproduce beyond its distributional boundary at similar rates than inside its range in N Spain. The seaweed was then expanding its distribution and the position of the boundary was set by dispersal limitations. At present, the southern boundary of this species seems to be directly influenced by very unfavourable abiotic conditions, which may be linked to the present scenario of climatic change or to environmental fluctuations acting at shorter‐time scales.

Summary 1The effects of density on the vital rates, growth form and population size structure of the invasive seaweed Sargassum muticum (Yendo) Fensholt were evaluated experimentally. 2 The initial difference in plant number between the highest and the lowest density (200 and 6400 plants m − 2 ) was considerably reduced by the end of the experiment. Surprisingly, this was mostly due to numbers increasing at lower densities, probably because microscopic forms were not removed during the experimental thinning. 3 The allometric length-dry mass relationship fitted a simple linear model on log-log scale for both the highest and the lowest densities, but had different slopes. At higher densities plants became taller and thinner as a consequence of variations in the production and growth of modules. 4 Mean size (dry mass) and the development of size hierarchies of plants were affected by both the addition of further (microscopic) recruits and asymmetrical competition among plants. Plant length distributions were also influenced by changes in the growth form of plants. The length hierarchies of main branches also suggested the presence of asymmetrical competition at this modular level. 5 Density influenced both mean size and morphology of the plants and thus induced changes in reproduction. The negative effect of density on individual plant size reduced the percentage of fertile plants and possibly their annual reproductive allocation, but these effects may be attenuated by morphological responses. 6 The responses of S. muticum to crowding are closely linked to its ability to colonize bare space. The massive reproductive output and very limited dispersal range account for local and dense recruitment patterns. Our results suggest that the responses of S. muticum to crowding allow the establishment of dense populations with high persistence and resistance to colonization by other species.

Anthropogenically induced global climate change has important implications for marine ecosystems with unprecedented ecological and economic consequences. Climate change will include the simultaneous increase of temperature and CO2 concentration in oceans. However, experimental manipulations of these factors at the community scale are rare. In this study, we used an experimental approach in mesocosms to analyse the combined effects of elevated CO2 and temperature on macroalgal assemblages from intertidal rock pools. Our model systems were synthetic assemblages of varying diversity and understory component and canopy species identity. We used assemblages invaded by the non‐indigenous canopy forming alga Sargassum muticum and assemblages with the native canopy species Cystoseira tamariscifolia. We examined the effects of both climate change factors on several ecosystem functioning variables (i.e. photosynthetic efficiency, productivity, respiration and biomass) and how these effects could be shaped by the diversity and species identity of assemblages. CO2 alone or in combination with temperature affected the performance of macroalgae at both individual and assemblage level. In particular, high CO2 and high temperature (20°C) drastically reduced the biomass of macroalgal assemblages and affected their productivity and respiration rates. The identity of canopy species also played an important role in shaping assemblage responses, whereas species richness did not seem to affect such responses. Species belonging to the same functional effect group responded differently to the same environmental conditions. Data suggested that assemblages invaded with S. muticum might be more resistant in a future scenario of climate change. Thus, in a future scenario of increasing temperature and CO2 concentration, macroalgal assemblages invaded with canopy‐forming species sharing response traits similar to those of S. muticum could be favoured.