Nutrient enrichment stimulates herbivory and alters epibiont assemblages at the edge but not inside subtidal macroalgal forests

AbstractNutrient enrichment is a major threat to subtidal macroalgal forests. Several studies have shown that nutrient inputs can enhance the ability of opportunistic algal species to acquire space freed by disturbance, at the expense of architecturally complex species that form forests. However, competition between canopy- and turf-forming macroalgae is not limited to the aftermath of disturbance. Canopy-forming macroalgae can provide suitable substratum for diverse epibiont assemblages, including both algae (epiphytes) and sessile invertebrates (epizoans). Despite evidence of enhanced epiphyte loading under eutrophic conditions, few experimental studies have assessed how nutrient enrichment influences the structure of epibiont assemblages on canopy-forming macroalgae at the edge versus inside forests. In oligotrophic waters of the NW Mediterranean, we experimentally tested the hypothesis that nutrient-driven proliferation of opportunistic epiphytic algae would affect the performance of the fucoid, Carpodesmia brachycarpa, and reduce the richness and abundance of the epizoan species they support. We predicted negative effects of nutrient enrichment to be greater at the edge than inside forests and on thalli that had recovered in cleared areas than on those within undisturbed canopy stands. Nutrient enrichment did not affect the photosynthetic efficiency and reproductive output of C. brachycarpa. By contrast, it enhanced herbivore consumption and decreased the cover and diversity of epizoans at forest edges, likely by stimulating the foraging activity of Arbacia lixula, the most abundant sea urchin in adjacent encrusting coralline barrens. Fertilization of areas inside forests had no effect on either C. brachycarpa or epibiont assemblages. Finally, nutrient enrichment effects did not vary between cleared and undisturbed areas. Our results show that moderate nutrient enrichment of oligotrophic waters does not necessarily cause the proliferation of epiphytes and, hence, a strengthening of their competitive effects on canopy-forming macroalgae. Nevertheless, enhanced herbivory damage to fertilized thalli at forest edges suggests that fragmentation could reduce the resilience of macroalgal forests and associated epibiont assemblages to nutrient enrichment

Propagules are not all equal: traits of vegetative fragments and disturbance regulate invasion success

Invasion success is regulated by multiple factors. While the roles of disturbance and propagule pressure in regulating the establishment of non-native species are widely acknowledged, that of propagule morphology (a proxy for quality) is poorly known. By means of a multi-factorial field experiment, we tested how the number (5 versus 10) and quality (intact, without fronds or without rhizoids) of fragments of the clonal invasive seaweed, Caulerpa cylindracea, influenced its ability to establish in patches of the native seagrass, Posidonia oceanica, exposed to different intensities of disturbance (0, 50 or 100% reduction in canopy cover). We hypothesized that the ability of fragments to establish would be greater for intact fragments (high quality) and reduced more by frond removal (low quality) than rhizoid removal (intermediate quality). At low propagule pressure or quality, fragment establishment was predicted to increase with increasing disturbance, whereas, at high propagule pressure or quality, it was predicted to be high regardless of disturbance intensity. Disturbance intensity, fragment number and quality had independent effects on C. cylindracea establishment success. Disturbance always facilitated fragment establishment. However, fragments retaining fronds, either intact or deprived of rhizoids, had higher establishment success than fragments deprived of fronds. Increasing propagule number had weak effects on the cover of C. cylindracea. Our results demonstrate that propagule traits enabling the acquisition of resources made available by disturbance can be more important than propagule number in determining the establishment and spread of clonal non-native plants. More generally, our study suggests that propagule quality is a key, yet underexplored, determinant of invasion success

Avoiding “Paper Parks”: A Global Literature Review on Socioeconomic Factors Underpinning the Effectiveness of Marine Protected Areas

Marine protected areas (MPAs) are a common management tool for preserving marine biodiversity and halting resource depletion. Despite the number of MPAs rapidly increasing worldwide, there are concerns over the full achievement of their objectives. Indeed, in some cases—the phenomenon of so-called “paper parks”—protected areas totally fail to achieve their conservation and socioeconomic targets. Therefore, identifying the factors underpinning MPA success or failure is crucial to increase their effectiveness. To achieve this goal, we performed a global literature review on the socioeconomic factors that managers should pursue to enhance MPA effectiveness on a global scale. A search of the Scopus database, using strings of keywords connected by Boolean operators, generated a batch of 715 items, out of which 68 were retained after the application of inclusion/exclusion criteria. Six other articles were added through the scanning of the literature cited in selected papers. We grouped MPA success-factors into 13 main groups and ranked them according to the frequency of citation in the literature. Our findings identify stakeholder involvement, increasing communication and awareness between specific stakeholder groups, as well as ensuring appropriate enforcement and monitoring, control and surveillance, as the leading factors for MPA success. Our results will assist in the process of upcoming global expansion of MPAs, thus contributing to improving conservation of marine biodiversity and associated livelihoods

Facilitation and the niche: Implications for coexistence, range shifts and ecosystem functioning

Viewing facilitation through the lens of the niche concept is one way to unify conceptual and empirical advances about the role of facilitation in community ecology. We clarify conceptually and through examples from marine and terrestrial environments how facilitation can expand species' niches and consider how these interactions can be scaled up to understand the importance of facilitation in setting a species' geographic range. We then integrate the niche-broadening influence of facilitation into current conceptual areas in ecology, including climate change, diversity maintenance and the relationship between diversity and ecosystem functioning. Because facilitation can influence the range of physical conditions under which a species can persist, it has the potential to mitigate the effects of climate change on species distributions. Whereas facilitation has mostly been considered as a diversity-promoting interaction by ameliorating abiotic stresses, if facilitated species' niches expand and become less distinct as a result of habitat amelioration, the forces that maintain diversity and promote coexistence in regions or habitats dominated by the facilitator could be reduced (i.e. the sign of the effects of facilitation on populations could be species-specific). Finally, shifting or broadening ecological niches could alter the relationship between diversity and ecosystem functioning. A niche-based perspective on the effects of facilitation can foster a greater mechanistic understanding of the role played by facilitation in regulating species coexistence, range shifts and ecosystem functioning in a changing world

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

Propagule composition regulates the success of an invasive seaweed across a heterogeneous seascape

Abstract Propagule pressure is acknowledged as a key determinant of invasion success. Nonetheless, the role of morphological or physiological attributes of propagules (i.e. their quality) in regulating invader establishment has been little explored. In particular, no study has investigated how the presence of propagules differing in quality within an inoculum influences establishment across heterogeneous landscapes. We experimentally tested the hypothesis that the quality (+Fronds+Rhizoids; +Fronds−Rhizoids; −Fronds+Rhizoids) and the diversity (1, 2 and 3 fragment types) of vegetative fragments of the seaweed Caulerpa taxifolia determine their establishment success across seascapes consisting of bare sediments and patches of the seagrass Zostera muelleri exposed to different disturbance intensities (control, seagrass canopy clipping and total removal). After 6 weeks, seaweed biomass, stolon and frond length, frond and rhizoid number were generally greater in unvegetated habitats (bare sediments and total seagrass removal) than full or reduced seagrass canopies. The type and the diversity of types of fragments inoculated had significant effects on the final biomass and morphological features of C. taxifolia only in vegetated habitats. In control plots, inocula of fragments retaining both fronds and rhizoids achieved higher biomass, developed longer stolons and more fronds. In canopy clipping plots, mixed inocula of +Fronds+Rhizoids and −Fronds+Rhizoids fragments had the greatest biomass and stolon length. Synthesis. Assessing how propagules differing in quality perform in different habitats might be not sufficient to draw a comprehensive picture of invasion risk, as their establishment can be modulated by both negative and positive interactions among them. Propagule composition should be, therefore, considered as a further dimension of propagule pressure. Our results also suggest that the relevance of specific propagule traits for invader establishment decreases from intact to degraded habitats. Considering propagule size in terms of amount of competent propagules, rather than an absolute measure, would refine our ability of predicting invasion risk across habitats differing in biotic or abiotic conditions

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

What are the effects of macroalgal blooms on the structure and functioning of marine ecosystems? A systematic review protocol

Abstract Background Anthropogenic activities are believed to have caused an increase in the magnitude, frequency, and extent of macroalgal blooms in marine and estuarine environments. These blooms may contribute to declines in seagrasses and non-blooming macroalgal beds, increasing hypoxia, and reductions in the diversity of benthic invertebrates. However, they may also provide other marine organisms with food and habitat, increase secondary production, and reduce eutrophication. The objective of this systematic review will be to quantify the positive and negative impacts of anthropogenically induced macroalgal blooms in order to determine their effects on ecosystem structure and functioning, and to identify factors that cause their effects to vary. Methods We will search a number of online databases to gather empirical evidence from the literature on the impacts of macroalgal blooms on: (1) species richness and other univariate measures of biodiversity; (2) productivity and abundance of algae, plants, and animals; and (3) biogeochemical cycling and other flows of energy and materials, including trophic interactions and cross-ecosystem subsidies. Data from relevant studies will be extracted and used in a random effects meta-analysis in order to estimate the average effect of macroalgal blooms on each response of interest. Where possible, sub-group analyses will be conducted in order to evaluate how the effects of macroalgal blooms vary according to: (1) which part of the ecosystem is being studied (e.g. which habitat type, taxonomic group, or trophic level); (2) the size of blooms; (3) the region in which blooms occurred; (4) background levels of ecosystem productivity; (5) physical and chemical conditions; (6) aspects of study design and quality (e.g. lab vs. field, experimental vs. observational, degree of replication); and (7) whether the blooms are believed to be anthropogenically induced or not

Investigating on the factors responsible for <i>Caulerpa racemosa</i> invasion = Indagini sui fattori responsabili dell'invasione di <i>Caulerpa racemosa</i>

The introduced alga Caulerpa racemosa (Forsskal) J. Agardh (Caulerpales, Chlorophyta) has become an important component of rocky assemblages in the subtidal of the Mediterranean. Understanding the faetors that regulate the establishment and spread of this species is, therefore, crucial to predicting future pathways of invasion and susceptible locales. Further, the aim of this study was to investigate on the factors responsible for Ihe successful invasion of C. racemosa in the Asinara Gulf (NW-Sardinia)