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Amphipods are one of the dominant epifaunal groups in seagrass meadows. However, our understanding of the biogeographical patterns in the distribution of these small crustaceans is limited. In this study, we investigated such patterns and the potential drivers in twelve Cymodocea nodosa meadows within four distinctive biogeographical areas across 2000 Km and 13° of latitude in two ocean basins (Mediterranean Sea and Atlantic Ocean). We found that species abundances in the assemblage of seagrass-associated amphipods differed among areas following a pattern largely explained by seagrass leaf area and epiphyte biomass, while the variation pattern in species presence/absence was determined by seagrass density and epiphyte biomass. Seagrass leaf area was also the most important determinant of greater amphipod total density and species richness, while amphipod density also increased with algal cover. Overall, our results evidenced that biogeographical patterns of variation in amphipod assemblages are mainly influenced by components of the habitat structure, which covary with environmental conditions, finding that structurally more complex meadows harboring higher abundance and richness of amphipods associated.

Despite the crucial role of herbivory in shaping community assembly, our understanding on biogeographical patterns of herbivory on seagrasses is limited compared to that on terrestrial plants. In particular, the drivers of such patterns remain largely unexplored. Here, we used a comparative-experimental approach in Cymodocea nodosa meadows, across all possible climate types within the seagrass distribution, 2000 km and 13° of latitude in two ocean basins, to investigate biogeographical variation in seagrass herbivory intensity and their drivers during July 2014. Particularly, the density and richness of herbivores and their food resources, seagrass size, carbon and nitrogen content, as well as latitude, sea surface temperature, salinity, chlorophyll, and sediment grain size, were tested as potential drivers. We found that shallow meadows can be subjected to intense herbivory, with variation in herbivory largely explained by fish density, seagrass size, and annual sea temperature range. The herbivorous fish density was the most important determinant of such variation, with the dominant seagrass consumer, the fish Sarpa salpa, absent at meadows from regions with low herbivory. In temperate regions where herbivorous fish are present, annual temperature ranges drive an intense summer herbivory, which is likely mediated not only by increased herbivore metabolic demands at higher temperatures, but also by higher fish densities. Invertebrate grazing (mainly by sea urchins, isopods, amphipods, and/or gastropods) was the dominant leaf herbivory in some temperate meadows, with grazing variation mainly influenced by seagrass shoot size. At the subtropical region (under reduced annual temperature range), lower shoot densities and seagrass nitrogen contents contributed to explain the almost null herbivory. We evidenced the combined influence of drivers acting at geographic (region) and local (meadow) scales, the understanding of which is critical for a clear prediction of variat

Despite the crucial role of herbivory in shaping community assembly, our understanding on biogeographical patterns of herbivory on seagrasses is limited compared to that on terrestrial plants. In particular, the drivers of such patterns remain largely unexplored. Here, we used a comparative-experimental approach in Cymodocea nodosa meadows, across all possible climate types within the seagrass distribution, 2000 km and 13° of latitude in two ocean basins, to investigate biogeographical variation in seagrass herbivory intensity and their drivers during July 2014. Particularly, the density and richness of herbivores and their food resources, seagrass size, carbon and nitrogen content, as well as latitude, sea surface temperature, salinity, chlorophyll, and sediment grain size, were tested as potential drivers. We found that shallow meadows can be subjected to intense herbivory, with variation in herbivory largely explained by fish density, seagrass size, and annual sea temperature range. The herbivorous fish density was themost important determinant of such variation,with the dominant seagrass consumer, the fish Sarpa salpa, absent atmeadows from regionswith lowherbivory. In temperate regionswhere herbivorous fish are present, annual temperature ranges drive an intense summer herbivory, which is likelymediated not only by increased herbivore metabolic demands at higher temperatures, but also by higher fish densities. Invertebrate grazing (mainly by sea urchins, isopods, amphipods, and/or gastropods) was the dominant leaf herbivory in some temperate meadows, with grazing variation mainly influenced by seagrass shoot size. At the subtropical region (under reduced annual temperature range), lower shoot densities and seagrass nitrogen contents contributed to explain the almost null herbivory.We evidenced the combined influence of drivers acting at geographic (region) and local (meadow) scales, the understanding of which is critical for a clear prediction of variation in s

To manage trace metal pollution it is critical to determine how much temporal trends can be attributed to local or large-scale sources. We tracked changes in metal content in the seagrass Posidonia oceanica, along the NW Mediterranean from 2003 to 2010. While Cu, Cd and Ni showed a large inter-site variation, likely due to local factors, Fe, Mn and Pb showed little local variation and synchronous interannual variability across sites, most likely due to large-scale sources. Zn showed equal importance of local and large-scale sources of variation. Temporal trends of Ni, Zn, Cd, Cu remained almost stable. In contrast, Fe, Mn and Pb slightly increased in the last decade. These trends suggest that metals like Cu, Cd, Ni can be effectively managed at local scale. Whereas, elements like Fe, Mn and Pb have an important large-scale component that needs to be managed across the frontiers of national jurisdictions.

Understanding how intra-specific differences in plant traits mediate vulnerability to herbivores of relevant habitat-forming plants is vital to attain a better knowledge on the drivers of the structure and functioning of ecosystems. Such studies, however, are rare in seagrass-mesograzer systems despite the increasingly recognized relevance of mesograzers as seagrass consumers. We investigated the role and potential trade-offs of multiple leaf traits in mediating the vulnerability of the seagrass Zostera noltei to different mesograzer species, the amphipod Gammarus insensibilis and the isopod Idotea chelipes. We worked with plants from two different meadows for which contrasting chemical and structural traits were expected based on previous information. We found that plants with high vulnerability to mesograzers (i.e. those preferred and subjected to higher rates of leaf area loss) had not only higher nitrogen content and lower C:N, fibre, and phenolics, but also tender and thinner leaves. No trade-offs between chemical and structural traits of the seagrass were detected, as they were positively correlated. When leaf physical structure was removed using agarreconstituted food, amphipod preference towards high-susceptibility plants disappeared; thus indicating that structural rather than chemical traits mediated the feeding preference. Removal of plant structure reduced the size of isopod preference to less than half, indicating a stronger contribution of structural traits (> 50%) but combined with chemical/nutritional traits in mediating the preference. We then hypothesized that the high environmental nutrient levels recorded in the meadow exhibiting high susceptibility modulate the differences observed between meadows in seagrass traits. To test this hypothesis, we exposed low-vulnerability shoots to eutrophic nutrient levels in a 6-week enrichment experiment. Nutrient enrichment increased Z. noltei nitrogen content and lowered C:N, fibre, and phenolics, but had no

Grazing-induced plant defences that reduce palatability to herbivores are widespread in terrestrial plants and seaweeds, but they have not yet been reported in seagrasses. We investigated the ability of two seagrass species to induce defences in response to direct grazing by three associated mesograzers. Specifically, we conducted feeding-assayed induction experiments to examine how mesograzer-specific grazing impact affects seagrass induction of defences within the context of the optimal defence theory. We found that the amphipod Gammarus insensibilis and the isopod Idotea chelipes exerted a low-intensity grazing on older blades of the seagrass Cymodocea nodosa, which reflects a weak grazing impact that may explain the lack of inducible defences. The isopod Synischia hectica exerted the strongest grazing impact on C. nodosa via high-intensity feeding on young blades with a higher fitness value. This isopod grazing induced defences in C. nodosa as indicated by a consistently lower consumption of blades previously grazed for 5, 12 and 16 days. The lower consumption was maintained when offered tissues with no plant structure (agarreconstituted food), but showing a reduced size of the previous grazing effect. This indicates that structural traits act in combination with chemical traits to reduce seagrass palatability to the isopod. Increase in total phenolics but not in C:N ratio and total nitrogen of grazed C. nodosa suggests chemical defences rather than a modified nutritional quality as primarily induced chemical traits. We detected no induction of defences in Zostera noltei, which showed the ability to replace moderate losses of young biomass to mesograzers via compensatory growth. Our study provides the first experimental evidence of induction of defences against meso-herbivory that reduce further consumption in seagrasses. It also emphasizes the relevance of grazer identity in determining the level of grazing impact triggering resistance and compensatory response

Coastal communities are under threat from many and often co-occurring local (e.g., pollution, eutrophication) and global stressors (e.g., climate change), yet understanding the interactive and cumulative impacts of multiple stressors in ecosystem function is far from being accomplished. Ecological redundancy may be key for ecosystem resilience, but there are still many gaps in our understanding of interspecific differences within a functional group, particularly regarding response diversity, that is, whether members of a functional group respond equally or differently to anthropogenic stressors. Herbivores are critical in determining plant community structure and the transfer of energy up the food web. Human disturbances may alter the ecological role of herbivory by modifying the defense strategies of plants and thus the feeding patterns and performance of herbivores. We conducted a suite of experiments to examine the independent and interactive effects of anthropogenic (nutrient and CO2 additions) and natural (simulated herbivory) disturbances on a seagrass and its interaction with two common generalist consumers to understand how multiple disturbances can impact both a foundation species and a key ecological function (herbivory) and to assess the potential existence of response diversity to anthropogenic and natural changes in these systems. While all three disturbances modified seagrass defense traits, there were contrasting responses of herbivores to such plant changes. Both CO2 and nutrient additions influenced herbivore feeding behavior, yet while sea urchins preferred nutrient-enriched seagrass tissue (regardless of other experimental treatments), isopods were deterred by these same plant tissues. In contrast, carbon enrichment deterred sea urchins and attracted isopods, while simulated herbivory only influenced isopod feeding choice. These contrasting responses of herbivores to disturbance-induced changes in seagrass help to better understand the ecological

Evaluating the efficacy of management actions to improve environmental quality is often difficult becausethere may be considerable lags before ecosystem management actions translate into measurable indi-cator responses. These delays make it difficult to justify often-expensive remedial actions to preventeutrophication. Therefore, it is critical to identify reliable, rapid and sensitive indicators to detect degra-dation and environmental quality improvement. We evaluate the efficacy of a set of indicators basedon the seagrass Posidonia oceanica to reliably and quickly detect ecosystem improvements using a 7-year (2003–2010) dataset of 10 stations along the Catalan coast (north-western Mediterranean Sea). Inthe Catalan region, environmental agencies have invested heavily on wastewater treatment, resultingin significant reductions (ca. 75%) in the BOD5discharged to coastal waters from 2003 to 2010. Theseimprovements were clearly reflected at the regional level (i.e. for all the stations averaged) in six bio-chemical seagrass indicators from our dataset. These indicators were directly related to eutrophication(nitrogen, 15N, phosphorus and total non-structural carbohydrates content in rhizomes, 34S and 13C inseagrass rhizomes and N content in epiphytes). In contrast, seagrass structural indicators, related to sea-grass abundance or meadow structure (density, cover) did not show any sign of overall recovery duringthe monitored period. These results confirm that biochemical seagrass indicators are the most sensitiveto water quality improvements within management time-scales (7–10 years) for slow-growing specieslike P. oceanica. Given the budgetary restrictions under which most management actions operate, theavailability of decision-support tools that function at appropriate time-scales is crucial to help managersvalidate the relative success of their remedial efforts. Our results indicate that low inertia, biochemicalseagrass indicators fit this task, and can be a robust

The pervasive use of ecological indices is increasingly requiring actions of harmonisation. Specifically, within the EU Water Framework Directive, an important effort in methods intercalibration is being done. However, a significant limitation in comparability assessment arises from the datasets used, which have different geographic origins. The purpose of our study was to perform an experimental intercalibration, where data were collected specifically on a set of common sites and following all the requirements of the methods being assessed. Three indices based on the marine angiosperm Posidonia oceanica, the POMI, the BiPo and the PoSte, were applied to sites in three different geographical areas of the western Mediterranean: Catalonia, Corsica and Southern Italy (Ischia), distant between hundreds and more than thousands of kilometers. Two indices, POMI and BiPo, showed not only a very good relationship with human pressures (measured on a common scale for all sites) but also a high comparability, in all aspects investigated. The differences found for the third one (PoSte) are hypothesised as being due to a different rationale used to define reference conditions, the different metrics used in the index, and in particular to a different definition of ecological status in relation to the time scale of the response to anthropogenic pressures. Our study demonstrates that indices with very different approaches can provide fully reliable and comparable results.

Biotic indices have become key assessment tools in most recent national and trans-national policies aimed at improving the quality of coastal waters and the integrity of their associated ecosystems. In this study we analyzed 90 published biotic indices, classified them into four types, and analyzed the strengths and weaknesses of each type in relation to the requirements of these policies. We identified three main type-specific weaknesses. First, the problems of applicability, due to practical and conceptual difficulties, which affect most indices related to ecosystem function. Second, the failure of many indices based on structural attributes of the community (e.g. taxonomic composition) to link deterioration with causative stressors, or to provide an early-detection capacity. Third, the poor relevance to the ecological integrity of indices based on attributes at the sub-individual level (e.g. multi-biomarkers). Additionally, most indices still fail on two further aspects: the broad-scale applicability and the definition of reference conditions. Nowadays, the most promising approach seems to be the aggregation of indices with complementary strengths, and obtained from different biological communities.

The influence of factors, both abiotic (light and nutrients) and biotic (meadow structure, herbivory and seagrass shoot length) in the abundance and distribution of epiphytes on the Mediterranean seagrass Posidonia oceanica was investigated by means of a correlational approach over a spatial scale of ca. 500 km and at two different depths (5 and 15 m). Variability was examined from the double perspective of integrative community measures (biomass, species richness and alpha-diversity) and species composition. We assessed the influence of biotic and abiotic factors on integrative community measures using multiple correlation analyses. The influence of these factors in the structure of the whole community was investigated using a distance-based RDA for a linear model. A total of 129 taxa, mostly epiphytic algae, were recorded across all sites and depths. A large part of the variability in species composition (51%) was explained by the variables investigated. In particular, variability caused by differences in grazing pressure was the most important (25%), followed by variables related to nutrient availability (11%), meadow structure (6%), light (5%) and seagrass shoot length (4%). Among integrative community variables investigated, species richness was also best explained by grazing and nutrients.

Eutrophication is one of the most relevant man-induced changes occurring in coastal waters. The identification and assessment of specific responses to eutrophication in seagrasses can provide a useful tool for the detection of changes in the water quality in coastal zones, given the wide range of distribution of these organisms. In this study, we combine a correlational (across-sites comparison) and a manipulative (fertilization experiment) approach to evaluate the usefulness and potential of alkaline phosphatase activity (APA) in the endemic Mediterranean seagrass Posidonia oceanica as an eutrophication biomarker. Our results showed that APA decreases promptly following nutrient additions, the response being maintained except during the winter period. APA also varies across natural meadows under different levels of nutrient discharges at scales relevant for monitoring purposes. AP activity seems to be an optimal ‘physiological biomarker’ that responds promptly and reliably to a pulse of eutrophication exposure. However, other considerations, such as the seasonality (the response disappears in winter), suggest its use with some caution and, as far as possible, as a complement of other bio-indicators.