Francisco Arenas

A continental scale evaluation of the role of limpet grazing on rocky shores

It is critical for our knowledge of biodiversity and ecosystem processes to understand how individual species contribute to ecosystem processes and how these contributions vary in space and time. We used a manipulative field experiment in five locations over 17° of latitude [from southern Portugal to the Isle of Man (British Isles)] to determine the relative response of rocky intertidal algal assemblages released from control by the grazing of limpets. Response ratios showed that when limpets were removed there was a trend of effects from north to south. In the north, grazing had a strong effect on algal assemblages, but removing grazers reduced spatial variability in assemblages. In the south, the effect of limpet grazing was far weaker and removal of grazers had a much reduced impact on spatial variability. Here we show a clear trophic control of an ecosystem in that grazing by limpets not only determines macroalgal abundance overall but also modifies ecosystem stability via variability in cover of algae.

THE INVASIBILITY OF MARINE ALGAL ASSEMBLAGES: ROLE OF FUNCTIONAL DIVERSITY AND IDENTITY

The emergence of the biodiversity-ecosystem functioning debate in the last decade has renewed interest in understanding why some communities are more easily invaded than others and how the impact of invasion on recipient communities and ecosystems varies. To date most of the research on invasibility has focused on taxonomic diversity, i.e., species richness. However, functional diversity of the communities should be more relevant for the resistance of the community to invasions, as the extent of functional differences among the species in an assemblage is a major determinant of ecosystem processes. Although coastal marine habitats are among the most heavily invaded ecosystems, studies on community invasibility and vulnerability in these habitats are scarce. We carried out a manipulative field experiment in tide pools of the rocky intertidal to test the hypothesis that increasing functional richness reduces the susceptibility of macroalgal communities to invasion. We selected a priori four functional groups on the basis of previous knowledge of local species characteristics: encrusting, turf, subcanopy, and canopy species. Synthetic assemblages containing one, two, three, or four different functional groups of seaweeds were created, and invasion by native species was monitored over an eight-month period. Cover and resource availability in the assemblages with only one functional group showed different patterns in the use of space and light, confirming true functional differences among our groups. Experimental results showed that the identity of functional groups was more important than functional richness in determining the ability of macroalgal communities to resist invasion and that resistance to invasion was resource-mediated.

Alien species and other notable records from a rapid assessment survey of marinas on the south coast of England

Arenas, F., Bishop, J.D.D., Carlton, J.T., Dyrynda, P.J., Farnham, W.F., Gonzalez, D.J., Jacobs, M.W., Lambert, C., Lambert, G., Nielsen, S.E., Pederson, J.A., Porter, J.S., Ward, S., Wood, C.A. (2006). Alien species and other notable records from a rapid assessment survey of marinas on the south coast of England. Journal of the Marine Biological Association of the United Kingdom, 86, (6), 1329-1337. Sponsorship: National Science Foundation grant IOB 0407527; Esme Fairbairn Foundation ALIENS project

A meta-analysis of seaweed impacts on seagrasses: Generalities and knowledge gaps

Seagrasses are important habitat-formers and ecosystem engineers that are under threat from bloom-forming seaweeds. These seaweeds have been suggested to outcompete the seagrasses, particularly when facilitated by eutrophication, causing regime shifts where green meadows and clear waters are replaced with unstable sediments, turbid waters, hypoxia, and poor habitat conditions for fishes and invertebrates. Understanding the situations under which seaweeds impact seagrasses on local patch scales can help proactive management and prevent losses at greater scales. Here, we provide a quantitative review of available published manipulative experiments (all conducted at the patch-scale), to test which attributes of seaweeds and seagrasses (e.g., their abundances, sizes, morphology, taxonomy, attachment type, or origin) influence impacts. Weighted and unweighted meta-analyses (Hedges d metric) of 59 experiments showed generally high variability in attribute-impact relationships. Our main significant findings were that (a) abundant seaweeds had stronger negative impacts on seagrasses than sparse seaweeds, (b) unattached and epiphytic seaweeds had stronger impacts than ‘rooted’ seaweeds, and (c) small seagrass species were more susceptible than larger species. Findings (a) and (c) were rather intuitive. It was more surprising that ‘rooted’ seaweeds had comparatively small impacts, particularly given that this category included the infamous invasive Caulerpa species. This result may reflect that seaweed biomass and/or shading and metabolic by-products like anoxia and sulphides could be lower for rooted seaweeds. In conclusion, our results represent simple and robust first-order generalities about seaweed impacts on seagrasses. This review also documented a limited number of primary studies. We therefore identified major knowledge gaps that need to be addressed before general predictive models on seaweed-seagrass interactions can be build, in order to effectively protect seagrass habitats from detrimental competition from seaweeds.

SIZE STRUCTURE AND DYNAMICS IN A POPULATION OF SARGASSUM MUTICUM (PHAEOPHYCEAE)

Sargassum muticum (Yendo) Fensholt is an introduced brown seaweed with a very distinctive seasonal growth cycle on European shores. The present study links the dynamics of a population of S. muticum with the seasonal growth cycle of the species and the density‐dependent processes operating throughout this cycle. Results indicate that both growth cycle and intraspecific competition influenced the structure and population dynamics. Size inequality increased during the slow growth phase (autumn–winter) of the 2‐year study. Mechanisms generating inequality of size could be the existence of asymmetric competition and the inherent differences in growth rates between old (regenerated) and new thalli (recruits). Inequality of size distributions decreased progressively during the last months of the growth phase (spring–summer) and could be related to a process of self‐thinning. There was a negative biomass–density relationship (as a measure of biomass accumulation‐driven mortality) that confirms the importance of self‐thinning as a major demographic factor in the S. muticum population.

Physical factors driving intertidal macroalgae distribution: physiological stress of a dominant fucoid at its southern limit

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 (Fv/Fm), 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.

Ecology of Sargassum muticum (Phaeophyta) on the north coast of Spain: III. Reproductive ecology

On the north coast of Spain, Sargassum muticum is fertile between spring and summer. Maturation occurs gradually from April to September. Senescence occurs simultaneously with full maturity increasing from June to September. Differences between two consecutive years can be explained by changes in environmental factors, such as seawater temperature and nutrient concentrations. Assuming that receptacles are the only reproductive structures, standing reproductive allocation (standing R A) in the population varied between 20-24% and the maximum value recorded for a single plant was 55%. There was a positive relationship between standing RA and the size of the plant, but this allometric relationship was much clearer when the number of receptacles (NR) was plotted against the weight of the plant. This suggests that NR is fixed for a given plant size but that actual allocation (standing RA) varied among plants. Inter-annual differences between reproductive investment were related to environmental factors and other factors intrinsic to the population.

Large-scale variation in combined impacts of canopy loss and disturbance on community structure and ecosystem functioning

Ecosystems are under pressure from multiple human disturbances whose impact may vary depending on environmental context. We experimentally evaluated variation in the separate and combined effects of the loss of a key functional group (canopy algae) and physical disturbance on rocky shore ecosystems at nine locations across Europe. Multivariate community structure was initially affected (during the first three to six months) at six locations but after 18 months, effects were apparent at only three. Loss of canopy caused increases in cover of non-canopy algae in the three locations in southern Europe and decreases in some northern locations. Measures of ecosystem functioning (community respiration, gross primary productivity, net primary productivity) were affected by loss of canopy at five of the six locations for which data were available. Short-term effects on community respiration were widespread, but effects were rare after 18 months. Functional changes corresponded with changes in community structure and/or species richness at most locations and times sampled, but no single aspect of biodiversity was an effective predictor of longer-term functional changes. Most ecosystems studied were able to compensate in functional terms for impacts caused by indiscriminate physical disturbance. The only consistent effect of disturbance was to increase cover of non-canopy species. Loss of canopy algae temporarily reduced community resistance to disturbance at only two locations and at two locations actually increased resistance. Resistance to disturbance-induced changes in gross primary productivity was reduced by loss of canopy algae at four locations. Location-specific variation in the effects of the same stressors argues for flexible frameworks for the management of marine environments. These results also highlight the need to analyse how species loss and other stressors combine and interact in different environmental contexts.

Physiological response of fucoid algae to environmental stress: comparing range centre and southern populations

Climate change has led to alterations in assemblage composition. Species of temperate macroalgae at their southern limits in the Iberian Peninsula have shown shifts in geographical range and a decline in abundance ultimately related to climate, but with the proximate factors largely unknown. We performed manipulative experiments to compare physiological responses of Fucus vesiculosus and Fucus spiralis from Portugal and Wales (UK), representing, respectively, southern and central areas of their distribution, to different intensities of solar radiation and different air temperatures. Following exposure to stressful emerged conditions, Portuguese and Welsh individuals of both fucoid species showed increased frond temperature, high desiccation levels and reduced photophysiological performance that was evident even after a 16 h recovery period, with light and temperature acting in an additive, not an interactive, manner. The level of physiological decline was influenced by geographical origin of populations and species identity, with algae from the south and those living higher on the shore coping better with stressful conditions. The negative effect of summer conditions on photophysiology may contribute to changes in fucoid abundance and distribution in southern Europe. Our results emphasise how physiological performance of geographically distinct populations can differ, which is particularly relevant when predicting responses to climate change.

Distribution and population dynamics of the introduced seaweed Grateloupia turuturu (Halymeniaceae, Rhodophyta) along the Portuguese coast

Araújo R., Violante J., Pereira R., Abreu H., Arenas F. and Sousa-Pinto I. 2011. Distribution and population dynamics of the introduced seaweed Grateloupia turuturu (Halymeniaceae, Rhodophyta) along the Portuguese coast. Phycologia 50: 392–402. DOI: 10.2216/10-65.1 Despite the increasing number of coastal areas invaded over the last years by the introduced seaweed Grateloupia turuturu, studies on this species are scarce worldwide, and its invasive capability and future impacts remain unexplored. In the present study, we describe the geographical distribution of the species after a few years of colonization of the Portuguese continental coastline. Additionally, we examine the structure and dynamics of one of the first population of Grateloupia in our shores. Seasonal dynamics of standing biomass, density, size structure and inequality (size variability), together with the presence of fertile fronds were investigated during one year. Grateloupia was recorded in midintertidal pools at 11 of the 36 localities sampled, two of which presenting an elevated abundance of fronds. Fronds were present during the entire year, but their size (length and biomass), density and size inequality varied seasonally reaching highest values at the beginning of the summer. The annual cycle of the population studied included a slow-growth phase in winter, a fast-growth phase during spring-early summer when some density-dependent regulation appeared to occur, as well as a dieback phase at the end of summer. In terms of fertility, abundance of fronds bearing reproductive structures were seasonally variable although carposporophytic and tetrasporophytic fronds were recorded throughout the year. The results of this work highlight some of the biological characteristics of Grateloupia turuturu that may determine its invasive nature and its possible impact on native ecosystems.