on freshwater and marine macrophytes: a review and perspective.Aquatic Botany http://dx.doi.org/10. 1016/j.aquabot.2016.04.008 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. AbstractUntil the 1990s, herbivory on aquatic vascular plants was considered to be of minor importance, and the predominant view was that freshwater and marine macrophytes did not take part in the food web: their primary fate was the detritivorous pathway. In the last 25 years, a substantial body of evidence has developed that shows that herbivory is an important factor in the ecology of vascular macrophytes across freshwater and marine habitats. Herbivores remove on average 40-48% of plant biomass in freshwater and marine ecosystems, which is typically 5-10 times greater than reported for terrestrial ecosystems. This may be explained by the lower C:N stoichiometry found in submerged plants.Herbivores affect plant abundance and species composition by grazing and bioturbation and therewith alter the functioning of aquatic ecosystems, including biogeochemical cycling, carbon stocks and primary production, transport of nutrients and propagules across ecosystem boundaries, habitat for other organisms and the level of shoreline protection by macrophyte beds.With ongoing global environmental change, herbivore impacts are predicted to increase.There are pressing needs to improve our management of undesirable herbivore impacts on macrophytes (e.g. leading to an ecosystem collapse), and the conflicts between people 4 associated with the impacts of charismatic mega-herbivores. While simultaneously, the longterm future of maintaining both viable herbivore populations and plant beds should be addressed, as both belong in complete ecosystems and have co-evolved in these long before the increasing influence of man. Better integration of the freshwater, marine, and terrestrial herbivory literatures would greatly benefit future research efforts.

Herbivory is a fundamental process that controls primary producer abundance and regulates energy and nutrient flows to higher trophic levels. Despite the recent proliferation of small-scale studies on herbivore effects on aquatic plants, there remains limited understanding of the factors that control consumer regulation of vascular plants in aquatic ecosystems. Our current knowledge of the regulation of primary producers has hindered efforts to understand the structure and functioning of aquatic ecosystems, and to manage such ecosystems effectively. We conducted a global meta-analysis of the outcomes of plant-herbivore interactions using a data set comprised of 326 values from 163 studies, in order to test two mechanistic hypotheses: first, that greater negative changes in plant abundance would be associated with higher herbivore biomass densities; second, that the magnitude of changes in plant abundance would vary with herbivore taxonomic identity. We found evidence that plant abundance declined with increased herbivore density, with plants eliminated at high densities. Significant between-taxa differences in impact were detected, with insects associated with smaller reductions in plant abundance than all other taxa. Similarly, birds caused smaller reductions in plant abundance than echinoderms, fish, or molluscs. Furthermore, larger reductions in plant abundance were detected for fish relative to crustaceans. We found a positive relationship between herbivore species richness and change in plant abundance, with the strongest reductions in plant abundance reported for low herbivore species richness, suggesting that greater herbivore diversity may protect against large reductions in plant abundance. Finally, we found that herbivore-plant nativeness was a key factor affecting the magnitude of herbivore impacts on plant abundance across a wide range of species assemblages. Assemblages comprised of invasive herbivores and native plant assemblages were associated with greater reductions in plant abundance compared with invasive herbivores and invasive plants, native herbivores and invasive plants, native herbivores and mixed-nativeness plants, and native herbivores and native plants. By contrast, assemblages comprised of native herbivores and invasive plants were associated with lower reductions in plant abundance compared with both mixed-nativeness herbivores and native plants, and native herbivores and native plants. However, the effects of herbivore-plant nativeness on changes in plant abundance were reduced at high herbivore densities. Our mean reductions in aquatic plant abundance are greater than those reported in the literature for terrestrial plants, but lower than aquatic algae. Our findings highlight the need for a substantial shift in how biologists incorporate plant-herbivore interactions into theories of aquatic ecosystem structure and functioning. Currently, the failure to incorporate top-down effects continues to hinder our capacity to understand and manage the ecological dynamics of habitats ...

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previous studies have considered waterfowl species as homologous, despite large differences in body mass, our results suggest that species body mass is a key determinant of waterfowl impact on plant standing crop.To examine relative impacts of waterfowl groups based on species body mass, a measure of plant biomass reduction (R s ) per bird per hectare was calculated for each group. Comparison of R s values indicated some differences in impact between different waterfowl groups, with swans having a greater per capita impact than smaller-bodied waterfowl groups. We present evidence that this difference is linked to disparities in individual body size and associated differences in intake rates, diet composition and energy requirements.Future research priorities are proposed, particularly the need for experiments that quantify the importance of factors that determine the magnitude of waterfowl impacts on plant standing crop.

Understanding plant community responses to combinations of biotic and abiotic factors is critical for predicting ecosystem response to environmental change. However, studies of plant community regulation have seldom considered how responses to such factors vary with the different phases of the plant growth cycle. To address this deficit we studied an aquatic plant community in an ecosystem subject to gradients in mute swan (Cygnus olor) herbivory, riparian shading, water temperature and distance downstream of the river source. We quantified abundance, species richness, evenness, flowering and dominance in relation to biotic and abiotic factors during the growth-, peak-, and recession-phases of the plant growth cycle. We show that the relative importance of biotic and abiotic factors varied between plant community properties and between different phases of the plant growth cycle. Herbivory became more important during the later phases of peak abundance and recession due to an influx of swans from adjacent pasture fields. Shading by riparian vegetation also had a greater depressing effect on biomass in later seasons, probably due to increased leaf abundance reducing light intensity reaching the aquatic plants. The effect of temperature on community diversity varied between upstream and downstream sites by altering the relative competitiveness of species at these sites. These results highlight the importance of seasonal patterns in the regulation of plant community structure and function by multiple factors.

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