Effects of resource pulse magnitude on nutrient availability,productivity, stability,and food web dynamics in experimental aquatic ecosystems

Oceanographic processes play a key role in influencing the structure of the marine planktonic ecosystems. Taking advantage of the quasi-simultaneous collection of a large ichthyoplanktonic dataset in different regions of the Central Mediterranean Sea (Italian/Maltese, Tunisian and Libyan waters), this study aimed at the identification of the main environmental drivers that control the structure of the larval fish assemblages. Spatial distribution and taxa composition were related to physical forcings (geostrophic currents and wind stress) and environmental conditions (bottom depth, temperature, salinity, chlorophyll-a concentration). ANOSIM and SIMPER identified contribution of fish taxa to the average Bray–Curtis dissimilarity among regions. In Italian and Libyan waters, two assemblages (neritic and oceanic) were identified, while a mixed assemblage characterized only some stations. Two neritic and one oceanic assemblages were discriminated in Tunisian waters. Random Forest classification model highlighted the essential role of the bathymetry, while Lagrangian simulations evidenced the action of the hydrodynamics in mixing neritic and oceanic assemblages in the Italian/Maltese and partially in Libyan waters. These findings highlighted the importance of the multidisciplinary approach and shed light on the potential value of the ichthyoplanktonic surveys for the assessment of the state of the marine ecosystem and the conservation of the fishery resources.     

Fish Assemblages of Mediterranean Marine Caves

Fish assemblages associated with 14 marine caves and adjacent external rocky reefs were investigated at four Marine Protected Areas (MPAs) along the coasts of Italy. Within the caves sampling was carried out in different sub-habitats: walls, ceilings, bottoms and ends of caves. On the whole, 38 species were recorded inside the 14 caves investigated. Eighteen species were exclusively found inside the caves: they were mainly represented by speleophilic (i.e. species preferentially or exclusively inhabiting caves) gobids (e.g. Didogobius splechtnai) and nocturnal species (e.g. Conger conger). Forty-one species were censused outside, 20 of which were shared with cave habitats. Apogon imberbis was the most common fish found in all 14 caves investigated, followed by Thorogobius ephippiatus (recorded in 13 caves), and Diplodus vulgaris and Scorpaena notata (both censused in 12 caves). Distinct fish assemblages were found between external rocky reefs and the different cave sub-habitats. New data on the distribution of some speleophilic gobids were collected, showing the existence of a pool of species shared by marine caves on a large scale (i.e. hundreds of km). Considering the uniqueness of cave fishes (18 exclusive species and different assemblage structures), the inclusion of marine caves among the habitats routinely investigated for fish biodiversity monitoring could facilitate the achievement of more comprehensive inventories. Due to their contribution to local species diversity and the shelter they provide to species valuable for conservation, marine caves should be prioritized for their inclusion not only within future MPAs through the Mediterranean Sea, but also into larger management spatial planning.     

Management of Local Stressors Can Improve the Resilience of Marine Canopy Algae to Global Stressors

Coastal systems are increasingly threatened by multiple local anthropogenic and global climatic stressors. With the difficulties in remediating global stressors, management requires alternative approaches that focus on local scales. We used manipulative experiments to test whether reducing local stressors (sediment load and nutrient concentrations) can improve the resilience of foundation species (canopy algae along temperate rocky coastlines) to future projected global climate stressors (high wave exposure, increasing sea surface temperature), which are less amenable to management actions. We focused on Fucoids (Cystoseira barbata) along the north-western Adriatic coast in the Mediterranean Sea because of their ecological relevance, sensitivity to a variety of human impacts, and declared conservation priority. At current levels of sediment and nutrients, C. barbata showed negative responses to the simulated future scenarios of high wave exposure and increased sea surface temperature. However, reducing the sediment load increased the survival of C. barbata recruits by 90.24% at high wave exposure while reducing nutrient concentrations resulted in a 20.14% increase in the survival and enhanced the growth of recruited juveniles at high temperature. We conclude that improving water quality by reducing nutrient concentrations, and particularly the sediment load, would significantly increase the resilience of C. barbata populations to projected increases in climate stressors. Developing and applying appropriate targets for specific local anthropogenic stressors could be an effective management action to halt the severe and ongoing loss of key marine habitats.     

A hazardous place to live: spatial and temporal patterns of species introduction in a hot spot of biological invasions

Globally, the spread of non-indigenous species in marine ecosystems is a major ecological and socio-economical concern. The need for long-term assessment on a large scale is a pre-requisite for understanding the drivers associated with their establishment and expansion. Here, the patterns of invasions of subtidal soft-bottom assemblages of shelled molluscs have been quantified based on a unique dataset collected between 2005 and 2012 along the coast of Israel (SE Mediterranean Sea), a hotspot of bioinvasion. Overall, the number of non-indigenous species doubled between 2005 and 2012. Significant differences in terms of species richness and relative abundance were observed in space and time in both native and non-indigenous species. A combination of enduring disturbance regimes related to human activities and site-specific environmental conditions seem to have a critical role in promoting the observed patterns. Our results emphasize the value of long term broad-scale systematic surveys to the development of effective environmental policies for the control of bioinvasions.     

Ecological effects of non‐native species in marine ecosystems relate to co‐occurring anthropogenic pressures

Predictors for the ecological effects of non‐native species are lacking, even though such knowledge is fundamental to manage non‐native species and mitigate their impacts. Current theories suggest that the ecological effects of non‐native species may be related to other concomitant anthropogenic stressors, but this has not been tested at a global scale. We combine an exhaustive meta‐analysis of the ecological effects of marine non‐native species with human footprint proxies to determine whether the ecological changes due to non‐native species are modulated by co‐occurring anthropogenic impacts. We found that non‐native species had greater negative effects on native biodiversity where human population was high and caused reductions in individual performance where cumulative human impacts were large. On this basis we identified several marine ecoregions where non‐native species may have the greatest ecological effects, including areas in the Mediterranean Sea and along the northwest coast of the United States. In conclusion, our global assessment suggests coexisting anthropogenic impacts can intensify the ecological effects of non‐native species.     

From global to local genetic structuring in the red gorgonian Paramuricea clavata: the interplay between oceanographic conditions and limited larval dispersal

Defining the scale of connectivity among marine populations and identifying the barriers to gene flow are tasks of fundamental importance for understanding the genetic structure of populations and for the design of marine reserves. Here, we investigated the population genetic structure at three spatial scales of the red gorgonian Paramuricea clavata (Cnidaria, Octocorallia), a key species dwelling in the coralligenous assemblages of the Mediterranean Sea. Colonies of P. clavata were collected from 39 locations across the Mediterranean Sea from Morocco to Turkey and analysed using microsatellite loci. Within three regions (Medes, Marseille and North Corsica), sampling was obtained from multiple locations and at different depths. Three different approaches (measures of genetic differentiation, Bayesian clustering and spatially explicit maximum‐difference algorithm) were used to determine the pattern of genetic structure. We identified genetic breaks in the spatial distribution of genetic diversity, which were concordant with oceanographic conditions in the Mediterranean Sea. We revealed a high level of genetic differentiation among populations and a pattern of isolation by distance across the studied area and within the three regions, underlining short effective larval dispersal in this species. We observed genetic differentiation among populations in the same locality dwelling at different depths, which may be explained by local oceanographic conditions and which may allow a process of local adaptation of the populations to their environment. We discuss the implications of our results for the conservation of the species, which is exposed to various threats.     

Distribution, ecology and morphology of Lytocarpia myriophyllum (Cnidaria: Hydrozoa), a Mediterranean Sea habitat former to protect

In the Mediterranean Sea, as well as in other parts of the word, intense bottom trawling threatens deep and mesophotic assemblages, compromising mainly the survivorship of erect organisms and of the habitat complexity they shape. Protection of species able to affect their habitats, by increasing spatial complexity and enhancing interspecific interactions, is crucial for biodiversity conservation. It is urgent to highlight the occurrence of those species which act as ecosystem engineers and/or habitat former to enhance awareness on their ecological role and to develop focused conservation strategies. Lytocarpia myriophyllum is the largest Leptomedusan hydroid of the Mediterranean Sea, with colonies up to 1 m high, and the most abundant Aglaopheniid in the eastern part of the North Atlantic Ocean. This species creates wide forests on soft bottoms stabilizing sediments, providing refuge and food for several other associated organisms and could be defined both a habitat former and an ecosystem engineer. Thanks to trimix diving here we report on new insights on the morphological, biological and ecological features of L. myriophyllum meadows from the Mediterranean Sea furnishing a baseline for protection plans focused on these facies. This work demonstrates that direct studies of mesophotic habitats allow to collect far more detailed information than grabs, ROVs, or towed camera arrays and highlights the urgent need to redefine the vertical extension of several marine protected areas.     

Habitat conservation prioritization: A floristic approach applied to a Mediterranean wetland network

The importance of plant communities for conservation purposes is recognized in their inclusion in the “Habitat Directive” (92/43 EEC), that relates habitat types to plant community syntaxonomic units. However, habitat definitions in the Habitat Directive lead to several inaccuracies in local habitat characterization. Several wetland plant communities (and their corresponding habitats), rare in the Mediterranean basin, are not included in the Habitat Directive. This study proposes criteria for assessing the conservation importance of habitats. It deals with plant community types at the alliance level, as promising units for setting conservation priorities. The principle criteria considered as drivers for setting alliance conservation values are the distribution and abundance of plant species of conservation interest and their fidelity to a plant community. Multivariate methods were used, and a quantitative floristic index of alliance conservation priority was created. This procedure was applied to an important wetland in central Italy. Results emphasize: (i) high conservation values of some alliances not listed in the Habitat Directive, confirming various gaps in the current conservation law affecting Mediterranean wetlands; (ii) that habitats widely distributed in other biogeographical areas, may greatly underestimate their conservation importance in Mediterranean region; (iii) need to consider regional peculiarities when setting conservation priorities.     

Facing complexity in tropical conservation: how reduced impact logging and climatic extremes affect beta diversity in tropical amphibian assemblages

Biodiversity in pristine forest biomes is increasingly disturbed by human activity. Drivers such as logging and climate extremes are thought to collectively erode diversity, but their interactions are not well understood. However, ignoring such complexities may result in poor conservation management decisions. Here, we present the first study dealing with the complexity arising from the effects of interactions of two increasingly important disturbance factors (selective logging and climatic extreme events) on beta diversity patterns at different scales. Specifically, we examined extensive amphibian assemblage datasets obtained within a quasi‐experimental pre‐/post‐harvesting scheme in the lowland rainforests of Central Guyana. Changes in small‐scale patterns of beta diversity were not detectable at the higher landscape level, indicating that local‐scale dynamics are more informative for evaluating disturbance impacts. The results also underscore the importance of including abundance data when investigating homogenization or heterogenization effects, which should be considered when designing post‐logging impact assessments and selecting impact indicators. Moreover, logging should be regarded as a multifaceted driver that contributes to changes in biodiversity patterns in different ways, depending on interactions with other drivers. The effects of extreme climate events were significantly more pronounced in unlogged forest, while logged forest assemblages appeared buffered due to the presence of novel habitats. Imprudent post‐logging renaturation measures may thus counteract conservation targets. These findings highlight the fact that indicator bias and unaccounted interactions between multiple drivers can lead to misguided management strategies.     

Mapping biodiversity in three-dimensions challenges marine conservation strategies: The example of coralligenous assemblages in North-Western Mediterranean Sea

Multi-facet diversity indices have been increasingly widely used in conservation ecology but congruence analyses both on horizontal and vertical axes have not yet been explored. We investigated the vertical and horizontal distributions of α and β taxonomic (TD), functional (FD) and phylogenetic diversity (PD) in a three-dimensional structured ecosystem. We focused on the Mediterranean coralligenous assemblages which form complex structures both vertically and horizontally, and are considered as the most diverse and threatened communities of the Mediterranean Sea. Although comparable to tropical reef assemblages in terms of richness, biomass and production, coralligenous assemblages are less known and more rarely studied, in particular because of their location in deep waters. Our study covers the entire range of distribution of coralligenous habitats along the French Mediterranean coasts, representing the most complete database so far developed for this important ecosystem. To our knowledge, this is the first analysis of spatial diversity patterns of marine biodiversity on both horizontal and vertical scales.Our study revealed that taxonomic diversity differed from functional and phylogenetic diversity patterns at the station level, the latter two being strongly structured by depth, with shallower stations generally richer than deeper ones. Considering all stations, phylogenetic diversity was less congruent to taxonomic diversity (Pearson's correlation of r = 0.48) but more congruent to functional diversity (r = 0.69) than randomly expected. Similar congruence patterns were revealed for stations deeper than 50 m (r = 0.44 and r = 0.84, respectively) but no significantly different congruence level than randomly expected was revealed among diversity facets for more shallow stations. Mean functional α- and β-diversity were lower than phylogenetic diversity and even lower than taxonomic α- and β-diversity for both vertical and horizontal scales. Low FD and PD values at both α- and β-diversity indicated functional and phylogenetic clustering. Community dissimilarities (β-diversity) increased over depth especially in central and eastern part of the French Mediterranean littoral and in northern Corsica, indicating coralligenous vertical structure within these regions. Overall horizontal β-diversity was higher within the 50–70 m depth belts.We conclude that taxonomic diversity alone is inadequate as a basis for setting conservation goals for this ecosystem and additional information, at least on phylogenetic diversity, is needed to preserve the ecosystem functioning and coralligenous evolutionary history. Our results highlight the necessity of considering different depth belts as a basis for regional scale conservation efforts. Current conservation approaches, such as the existing marine protected areas, are insufficient in preserving coralligenous habitats. The use of multi-facet indices should be considered, focusing on preserving local diversity patterns and compositional dissimilarities, both vertically and horizontally.     

Organization of Squamata (Reptilia) assemblages in Mediterranean archipelagos

Mediterranean islands have complex reptile assemblages, but little is known about the factors that determine their organization. In this study, the structure of assemblages of Squamata was evaluated based on their species richness and two measures of phylogenetic diversity (variability and clustering). I evaluated the composition of the assemblages comparing distinct biogeographic subregions within the Mediterranean: Adriatic, Aegean, Balearic, Corsica–Sardinia, Crete, Gulf of Gabés, Ionian Sea, Ligurian Sea, Malta, Sicily, and Tyrrhenian Sea. The effect of island environments and geographical isolation on the diversity metrics was assessed using generalized linear models. The analyses indicated that species richness was mostly influenced by island area and geographical isolation. Assemblages on smaller islands were poorer in species and phylogenetically dispersed, possibly as an effect of interspecific competition. The species composition of the assemblages was determined by similar environmental drivers within the biogeographic subregions, including island area, island elevation, geographical isolation, and aridity. In several subregions, significant patterns of phylogenetic attraction were found in species co‐occurrences, caused by the limits imposed by the island size on large predatory species.     

Macroalgal Composition Determines the Structure of Benthic Assemblages Colonizing Fragmented Habitats

Understanding the consequences of fragmentation of coastal habitats is an important topic of discussion in marine ecology. Research on the effects of fragmentation has revealed complex and context-dependent biotic responses, which prevent generalizations across different habitats or study organisms. The effects of fragmentation in marine environments have been rarely investigated across heterogeneous habitats, since most studies have focused on a single type of habitat or patch. In this study, we assessed the effects of different levels of fragmentation (i.e. decreasing size of patches without overall habitat loss). We measured these effects using assemblages of macro-invertebrates colonizing representative morphological groups of intertidal macroalgae (e.g. encrusting, turf and canopy-forming algae). For this purpose, we constructed artificial assemblages with different combinations of morphological groups and increasing levels of fragmentation by manipulating the amount of bare rock or the spatial arrangement of different species in mixed assemblages. In general, our results showed that 1) fragmentation did not significantly affect the assemblages of macroinvertebrates; 2) at greater levels of fragmentation, there were greater numbers of species in mixed algal assemblages, suggesting that higher habitat complexity promotes species colonization. Our results suggest that predicting the consequences of fragmentation in heterogeneous habitats is dependent on the type and diversity of morphological groups making up those habitats.     

Toward a conceptual framework for managing and conserving marine habitats: A case study of kelp forests in the Salish Sea

Kelp forests are in decline across much of their range due to place‐specific combinations of local and global stressors. Declines in kelp abundance can lead to cascading losses of biodiversity and productivity with far‐reaching ecological and socioeconomic consequences. The Salish Sea is a hotspot of kelp diversity where many species of kelp provide critical habitat and food for commercially, ecologically, and culturally important fish and invertebrate species. However, like other regions, kelp forests in much of the Salish Sea are in rapid decline. Data gaps and limited long‐term monitoring have hampered attempts to identify and manage for specific drivers of decline, despite the documented urgency to protect these important habitats. To address these knowledge gaps, we gathered a focus group of experts on kelp in the Salish Sea to identify perceived direct and indirect stressors facing kelp forests. We then conducted a comprehensive literature review of peer‐reviewed studies from the Salish Sea and temperate coastal ecosystems worldwide to assess the level of support for the pathways identified by the experts, and we identified knowledge gaps to prioritize future research. Our results revealed major research gaps within the Salish Sea and highlighted the potential to use expert knowledge for making informed decisions in the region. We found high support for the pathways in the global literature, with variable consensus on the relationship between stressors and responses across studies, confirming the influence of local ecological, oceanographic, and anthropogenic contexts and threshold effects on stressor–response relationships. Finally, we prioritized areas for future research in the Salish Sea. This study demonstrates the value expert opinion has to inform management decisions. These methods are readily adaptable to other ecosystem management contexts, and the results of this case study can be immediately applied to kelp management.     

Limited habitat and conservation value of a young artificial reef

Loss and degradation of natural habitats and their biodiversity may, arguably, be mitigated or compensated through the creation of human-engineered habitats: the underlying conservation tenet is that these artificial habitats compensate for diminished diversity caused by human impacts at local or regional scales. This approach is widely used in the sea by purposefully scuttling ships to create artificial reefs, but its performance as a conservation tool is seldom critically examined in these situations. Here we test if the diversity of sessile invertebrate assemblages on a large, but young (3 years), artificial reef, created by sinking a 133 m long battle ship off Eastern Australia, can mimic that of nearby natural reefs. We use this system as a model to test whether this artificial reef can form compensatory habitat of comparable quality and levels of biodiversity. Our assessment is based on the abundance, species richness, and species composition of sessile invertebrate assemblages, including corals. Despite some signs that temporal trajectories of ecological metrics, such as cover, began to approach natural conditions after 3 years, the ecological structure of sessile invertebrate assemblages on this young wreck remained fundamentally different from those on nearby natural reefs. In particular, large, long-lived corals were abundant on natural rocky reefs, but were rare and covered little area on the young wreck. These data demonstrate that when trajectories to community convergence with natural habitats are prolonged, as may be the case here, any compensatory effects of artificial habitats will have a considerable time lag. Such lags have implications for appraising the conservation value of wrecks and artificial reefs, and they emphasize the need to explicitly acknowledge temporal dynamics when using artificial habitats as complementary conservation tools to augment larger conservation efforts on natural systems.     

Synergistic effects of climate change and local stressors: CO2 and nutrient-driven change in subtidal rocky habitats

Climate-driven change represents the cumulative effect of global through local-scale conditions, and understanding their manifestation at local scales can empower local management. Change in the dominance of habitats is often the product of local nutrient pollution that occurs at relatively local scales (i.e. catchment scale), a critical scale of management at which global impacts will manifest. We tested whether forecasted global-scale change [elevated carbon dioxide (CO₂) and subsequent ocean acidification] and local stressors (elevated nutrients) can combine to accelerate the expansion of filamentous turfs at the expense of calcifying algae (kelp understorey). Our results not only support this model of future change, but also highlight the synergistic effects of future CO₂ and nutrient concentrations on the abundance of turfs. These results suggest that global and local stressors need to be assessed in meaningful combinations so that the anticipated effects of climate change do not create the false impression that, however complex, climate change will produce smaller effects than reality. These findings empower local managers because they show that policies of reducing local stressors (e.g. nutrient pollution) can reduce the effects of global stressors not under their governance (e.g. ocean acidification). The connection between research and government policy provides an example whereby knowledge (and decision making) across local through global scales provides solutions to some of the most vexing challenges for attaining social goals of sustainability, biological conservation and economic development.     

Protected and threatened components of fish biodiversity in the Mediterranean sea

The Mediterranean Sea (0.82% of the global oceanic surface) holds 4%-18% of all known marine species (~17,000), with a high proportion of endemism [1, 2]. This exceptional biodiversity is under severe threats [1] but benefits from a system of 100 marine protected areas (MPAs). Surprisingly, the spatial congruence of fish biodiversity hot spots with this MPA system and the areas of high fishing pressure has not been assessed. Moreover, evolutionary and functional breadth of species assemblages [3] has been largely overlooked in marine systems. Here we adopted a multifaceted approach to biodiversity by considering the species richness of total, endemic, and threatened coastal fish assemblages as well as their functional and phylogenetic diversity. We show that these fish biodiversity components are spatially mismatched. The MPA system covers a small surface of the Mediterranean (0.4%) and is spatially congruent with the hot spots of all taxonomic components of fish diversity. However, it misses hot spots of functional and phylogenetic diversity. In addition, hot spots of endemic species richness and phylogenetic diversity are spatially congruent with hot spots of fishery impact. Our results highlight that future conservation strategies and assessment efficiency of current reserve systems will need to be revisited after deconstructing the different components of biodiversity.     

Projected impacts of climate warming on the functional and phylogenetic components of coastal Mediterranean fish biodiversity

Climate warming affects biodiversity distribution across all ecosystems. However, beyond changes in species richness, impacts on other biodiversity components are still overlooked, particularly in the marine realm. Here we forecasted the potential effect of climate warming on the phylogenetic and functional components of coastal Mediterranean fish biodiversity. To do so, we used species distribution models to project the potential distribution of 230 coastal fish species by the end of the 21st century based on the IPCC A2 scenario implemented with the Mediterranean climatic model NEMOMED8. From these projections, we assessed the changes in phylogenetic (PD) and functional diversity (FD) of fish assemblages at multiple spatial scales using a dated molecular phylogeny and an extensive functional trait database. At the scale of the entire Mediterranean Sea, the projected extinctions of 40 coastal fish species would lead to a concomitant erosion of PD and FD (13.6 and 3%, respectively). However, a null model revealed that species loss at this scale would not lead to a disproportionate erosion of PD and FD. Similar results were found when considering fish assemblages at the grid cell scale. Indeed, at this scale, the projected changes in species richness would lead to unexpected losses of PD and FD for localized and small areas only. A disproportionate erosion of PD under climate warming was only forecasted when analysing fish assemblages at an intermediate spatial scale, namely the Mediterranean marine ecoregions. Overall, our results emphasize the importance of considering multiple spatial scales when assessing potential impacts of climate warming on the multiple components marine biodiversity.     

Methodological challenges in assessing the environmental status of a marine ecosystem: case study of the Baltic Sea

Assessments of the environmental status of marine ecosystems are increasingly needed to inform management decisions and regulate human pressures to meet the objectives of environmental policies. This paper addresses some generic methodological challenges and related uncertainties involved in marine ecosystem assessment, using the central Baltic Sea as a case study. The objectives of good environmental status of the Baltic Sea are largely focusing on biodiversity, eutrophication and hazardous substances. In this paper, we conduct comparative evaluations of the status of these three segments, by applying different methodological approaches. Our analyses indicate that the assessment results are sensitive to a selection of indicators for ecological quality objectives that are affected by a broad spectrum of human activities and natural processes (biodiversity), less so for objectives that are influenced by a relatively narrow array of drivers (eutrophications, hazardous substances). The choice of indicator aggregation rule appeared to be of essential importance for assessment results for all three segments, whereas the hierarchical structure of indicators had only a minor influence. Trend-based assessment was shown to be a useful supplement to reference-based evaluation, being independent of the problems related to defining reference values and indicator aggregation methodologies. Results of this study will help in setting priorities for future efforts to improve environmental assessments in the Baltic Sea and elsewhere, and to ensure the transparency of the assessment procedure.