Spatial correlation reverses the compound effect of multiple stressors on rocky shore biofilm

Understanding how multifactorial fluctuating environments affect species and communities remains one of the major challenges in ecology. The spatial configuration of the environment is known to generate complex patterns of correlation among multiple stressors. However, to what extent the spatial correlation between simultaneously fluctuating variables affects ecological assemblages in real-world conditions remains poorly understood. Here, we use field experiments and simulations to assess the influence of spatial correlation of two relevant climate variables - warming and sediment deposition following heavy precipitation - on the biomass and photosynthetic activity of rocky intertidal biofilm. First, we used a response-surface design experiment to establish the relation between biofilm, warming, and sediment deposition in the field. Second, we used the response surface to generate predictions of biofilm performance under different scenarios of warming and sediment correlation. Finally, we tested the predicted outcomes by manipulating the degree of correlation between the two climate variables in a second field experiment. Simulations stemming from the experimentally derived response surface showed how the degree and direction (positive or negative) of spatial correlation between warming and sediment deposition ultimately determined the nonlinear response of biofilm biomass (but not photosynthetic activity) to fluctuating levels of the two climate variables. Experimental results corroborated these predictions, probing the buffering effect of negative spatial correlation against extreme levels of warming and sediment deposition. Together, these results indicate that consideration of nonlinear response functions and local-scale patterns of correlation between climate drivers can improve our understanding and ability to predict ecological responses to multiple processes in heterogeneous environments

Connell and Slatyer's models of succession in the biodiversity era

Understanding how species interactions drive succession is a key issue in ecology. In this study we show the utility of combining the concepts and methodologies developed within the biodiversity–ecosystem functioning research program with J. H. Connell and R. O. Slatyer’s classic framework to understand succession in assemblages where multiple interactions between early and late colonists may include both inhibitory and facilitative effects. We assessed the net effect of multiple species interactions on successional changes by manipulating the richness, composition, and abundance of early colonists in a low-shore assemblage of algae and invertebrates of the northwestern Mediterranean. Results revealed how concomitant changes in species richness and abundance can strongly alter the net effect of inhibitory vs. facilitative interactions on succession. Increasing richness of early colonists inhibited succession, but only under high levels of initial abundance, probably reflecting the formation of a highly intricate matrix that prevented further colonization. In contrast, increasing initial abundance of early colonists tended to facilitate succession under low richness. Thus, changes in abundance of early colonists mediated the effects of richness on succession

Beyond Competition: Incorporating Positive Interactions between Species to Predict Ecosystem Invasibility

Incorporating positive species interactions into models relating native species richness to community invasibility will increase our ability to forecast, prevent, and manage future invasions

Subtle differences in growth rate drive contrasting responses of ephemeral primary producers to recurrent disturbances

Although the importance of time after disturbance is well established in the ecological literature, studies examining how differences in growth rate affect species recovery and persistence in relation to the interval between recurrent perturbations are rare. We examined the response of two ephemeral primary producers inhabiting high-shore rock pools, epilithic microphytobenthos (EMPB), and green filamentous algae, to disturbance regimes varying for the time interval between consecutive events. Informed from an empirically parametrized growth model's outcomes, we tested the hypothesis that EMPB would be able to recover from more frequent disturbance compared with filamentous algae in a field experiment involving three physical disturbance patterns differing for the clustering degree: high, moderate, and low (20, 40, and 80 days between disturbances). We predicted that: high clustering would prevent the recovery of both taxa; moderate clustering would prevent the recovery of the slower growing taxon only (filamentous algae); both taxa would recover under low clustering. Results showed that EMPB persisted independently of the clustering degree, whereas filamentous algae did not withstand any disturbance regime. Dramatically different effects of disturbance on organisms with subtle differences in their growth rate indicate that even stronger responses may be expected from taxa with more markedly contrasting life histories

Effects of grazer diversity on marine microphytobenthic biofilm: a ‘tug of war’ between complementarity and competition

Species loss is one of the most striking problems related to human-driven environmental changes. Nevertheless, biodiversity and ecosystem functioning experiments have mainly focused on primary producers, paying less attention to the consequences of changing diversity at higher trophic levels. We performed a field experiment using cage enclosures to test the effects of species richness, identity and density of gastropod grazers on the photosynthetic efficiency and biomass of intertidal biofilm on an exposed rocky shore in the northwest Mediterranean. The diversity and composition of intertidal grazers affected the photosynthetic efficiency of biofilm with only negligible effects on biomass. Individual species showed strong identity effects. In assemblages of 2 or more species, positive or negative complementarity effects occurred. The magnitude of the ecosystem response is expected to depend on the particular species assemblage and its density, which will determine whether niche partitioning or competition is the prevailing process. Grazer preference in specific components of biofilm, characterized by different photosynthetic efficiency and competitive abilities, might explain concomitant changes in photosynthetic efficiency and comparable levels in biomass among treatments. The effects of grazers declined following the natural trend of decreasing biomass of biofilm during the study period, highlighting the importance of considering temporal variability in the effects of biodiversity on ecosystem functioning. This work emphasizes the key role of species identity to predict effects on their resources and ecosystem functioning.This work was partially supported by the University of Pisa

Multifractal spatial distribution of epilithic microphytobenthos on a Mediterranean rocky shore

Understanding how patterns and processes relate across spatial scales is one of the major goals in ecology. 1/f models have been applied mostly to time series of environmental and ecological variables, but they can also be used to analyse spatial patterns. Since 1/f noise may display scale-invariant behaviour, ecological phenomena whose spatial variability shows 1/f type scaling are susceptible to further characterization using fractals or multifractals. Here we use spectral analysis and multifractal techniques (generalized dimension spectrum) to investigate the spatial distribution of epilithic microphytobenthos (EMPB) on rocky intertidal surfaces. EMPB biomass was estimated from calibrated colour-infrared images that provided indirect measures of rock surface chlorophyll a concentration, along two 8-m and one 4-m long transects sampled in January and November 2012. Results highlighted a pattern of spectral coefficient close to or greater than one for EMPB biomass distribution and multifractal structures, that were consistent among transects, implying scale-invariance in the spatial distribution of EMPB. These outcomes can be interpreted as a result of the superimposition of several biotic and abiotic processes acting at multiple spatial scales. However, the scale-invariant nature of EMPB spatial patterns can also be considered a hallmark of self-organization, underlying the possible role of scale-dependent feedback in shaping EMPB biomass distribution.This work is part of a requirement for a PhD by MDB and was partially supported by the Univ. of Pisa and by the FP 7 EU project VECTORS “VECTORS of Change in Oceans and Seas Marine Life, Impact on Economic Sectors”. CS was supported by a postdoctoral grant from the Alfonso Martin Escudero Foundation from Spain

The effects of exotic seaweeds on native benthic assemblages: variability between trophic levels and influence of background environmental and biological conditions

Abstract Background Biological invasions are among the most severe threats to marine biodiversity. The impacts of introduced seaweeds on native macroalgal assemblages have been thoroughly reviewed. In contrast, no attempt has been made to synthesize the available information on the effects of exotic seaweeds on other trophic levels. In addition, it has not been clarified whether the effects of introduced seaweeds on native assemblages vary according to background physical and biological conditions. Methods This protocol provides details of our proposed method to carry out a systematic review aiming to identify and synthesize existing knowledge to answer the following primary questions: a) how does the impact of the presence of exotic seaweeds on native primary consumers (across trophic levels) compare in magnitude and extent to that observed on native primary producers (same trophic level)?; b) does the intensity of the effects of the presence of exotic seaweeds on native benthic ecosystems vary along a gradient of human disturbance (i.e. from urban/industrial areas to extra-urban areas to pristine areas)

Unveiling the complexity and ecological function of aquatic macrophyte-animal networks in coastal ecosystems

Network theory offers innovative tools to explore the complex ecological mechanisms regulating species associations and interactions. Although interest in ecological networks has grown steadily during the last two decades, the application of network approaches has been unequally distributed across different study systems: while some kinds of interactions (e.g. plant–pollinator and host–parasite) have been extensively investigated, others remain relatively unexplored. Among the latter, aquatic macrophyte–animal associations in coastal environments have been largely neglected, despite their major role in littoral ecosystems. The ubiquity of macrophyte systems, their accessibility and multi-faceted ecological, economical and societal importance make macrophyte–animal systems an ideal subject for ecological network science. In fact, macrophyte–animal networks offer an aquatic counterpart to terrestrial plant–animal networks. In this review, we show how the application of network analysis to aquatic macrophyte–animal associations has the potential to broaden our understanding of how coastal ecosystems function. Network analysis can also provide a key to understanding how such ecosystems will respond to on-going and future threats from anthropogenic disturbance and environmental change. For this, we: (i) identify key issues that have limited the application of network theory and modelling to aquatic animal– macrophyte associations; (ii) illustrate through examples based on empirical data how network analysis can offer new insights on the complexity and functioning of coastal ecosystems; and (iii) provide suggestions for how to design future studies and establish this new research line into network ecology.Peer reviewe

Deterministic Factors Overwhelm Stochastic Environmental Fluctuations as Drivers of Jellyfish Outbreaks

16 pages, 4 figures, 1 table, supporting Information http://dx.doi.org/10.1371/journal.pone.0141060Jellyfish outbreaks are increasingly viewed as a deterministic response to escalating levels of environmental degradation and climate extremes. However, a comprehensive understanding of the influence of deterministic drivers and stochastic environmental variations favouring population renewal processes has remained elusive. This study quantifies the deterministic and stochastic components of environmental change that lead to outbreaks of the jellyfish Pelagia noctiluca in the Mediterranen Sea. Using data of jellyfish abundance collected at 241 sites along the Catalan coast from 2007 to 2010 we: (1) tested hypotheses about the influence of time-varying and spatial predictors of jellyfish outbreaks; (2) evaluated the relative importance of stochastic vs. deterministic forcing of outbreaks through the environmental bootstrap method; and (3) quantified return times of extreme events. Outbreaks were common in May and June and less likely in other summer months, which resulted in a negative relationship between outbreaks and SST. Cross- and along-shore advection by geostrophic flow were important concentrating forces of jellyfish, but most outbreaks occurred in the proximity of two canyons in the northern part of the study area. This result supported the recent hypothesis that canyons can funnel P. noctiluca blooms towards shore during upwelling. This can be a general, yet unappreciated mechanism leading to outbreaks of holoplanktonic jellyfish species. The environmental bootstrap indicated that stochastic environmental fluctuations have negligible effects on return times of outbreaks. Our analysis emphasized the importance of deterministic processes leading to jellyfish outbreaks compared to the stochastic component of environmental variation. A better understanding of how environmental drivers affect demographic and population processes in jellyfish species will increase the ability to anticipate jellyfish outbreaks in the futureThe authors gratefully acknowledge financial support by the European Community Seventh Framework Programme (FP7/2007–2013) for the project VECTORS (grant agreement no. 266445) (URL: http://cordis.europa.eu/fp7/home_en.html). AC was supported by a doctoral fellowship from the Chilean National Commission for Scientific and Technological Research (CONICYT – PFCHA/Doctorado al Extranjero 4a Convocatoria, 72120016).Peer Reviewe

The effects of an invasive seaweed on native communities vary along a gradient of land-based human impacts

The difficulty in teasing apart the effects of biological invasions from those of other anthropogenic perturbations has hampered our understanding of the mechanisms underpinning the global biodiversity crisis. The recent elaboration of global-scale maps of cumulative human impacts provides a unique opportunity to assess how the impact of invaders varies among areas exposed to different anthropogenic activities. A recent meta-analysis has shown that the effects of invasive seaweeds on native biota tend to be more negative in relatively pristine than in human-impacted environments. Here, we tested this hypothesis through the experimental removal of the invasive green seaweed, Caulerpa cylindracea, from rocky reefs across the Mediterranean Sea. More specifically, we assessed which out of land-based and sea-based cumulative impact scores was a better predictor of the direction and magnitude of the effects of this seaweed on extant and recovering native assemblages. Approximately 15 months after the start of the experiment, the removal of C. cylindracea from extant assemblages enhanced the cover of canopy-forming macroalgae at relatively pristine sites. This did not, however, result in major changes in total cover or species richness of native assemblages. Preventing C. cylindracea re-invasion of cleared plots at pristine sites promoted the recovery of canopyforming and encrusting macroalgae and hampered that of algal turfs, ultimately resulting in increased species richness. These effects weakened progressively with increasing levels of land-based human impacts and, indeed, shifted in sign at the upper end of the gradient investigated. Thus, at sites exposed to intense disturbance from land-based human activities, the removal of C. cylindracea fostered the cover of algal turfs and decreased that of encrusting algae, with no net effect on species richness. Our results suggests that competition from C. cylindracea is an important determinant of benthic assemblage diversity in pristine environments, but less so in species-poor assemblages found at sites exposed to intense disturbance from landbased human activities, where either adverse physical factors or lack of propagules may constrain the number of potential native colonizers. Implementing measures to reduce the establishment and spread of C. cylindracea in areas little impacted by land-based human activities should be considered a priority for preserving the biodiversity of Mediterranean shallow rocky reefs