Abstract1. Our knowledge of the functional role of large herbivores is rapidly expanding, and the impact of grazing on species coexistence and nonnative species expansion has been studied across ecosystems. However, experimental data on large grazer impacts on plant invasion in aquatic ecosystems are lacking.2. Since its introduction in 2002, the seagrass species Halophila stipulacea has rapidly expanded across the Eastern Caribbean, forming dense meadows in green turtle (Chelonia mydas)-foraging areas. We investigate the changes in seagrass species coexistence and the impacts of leaf grazing by green turtles on nonnative seagrass expansion in Lac Bay (Bonaire, Caribbean Netherlands).3. Green turtle grazing behaviour changed after the introduction of nonnative seagrass to Lac Bay in 2010. Field observations, together with time-lapse satellite images over the last four decades, showed initiation of new grazing patches (65 ha, an increase of 72%). The sharp border between grazed and ungrazed seagrass patches moved in the direction of shallower areas with native seagrass species that had previously been ungrazed. Green turtles deployed withFastloc-GPS transmitters confirmed high site fidelity to these newly cropped patches. In addition, cafeteria experiments indicated selective grazing by green turtles on native species. These native seagrass species had significantly higher nutritional values compared to the nonnative species. In parallel, exclosure experiments showed that nonnative seagrass expanded more rapidly in grazed canopies | INTRODUC TI ONLarge herbivores, whether aquatic or terrestrial, can have strong impacts on associated species and can be critical ecosystem engineers as they alter plant productivity, modify geomorphology, and influence nutrient cycling, habitat structure and plant coexistence (Bakker et al., 2016;Poore et al., 2012;Wood, Armstron, & Richards, 1967). Megaherbivores can impact plant species coexistence and species composition via multiple mechanisms. Preferential grazing on dominant plant species can promote species diversity by releasing competitors (Olff & Ritchie, 1998), or reduce diversity by selectively removing nondominant species (Hidding, Bakker, et al., 2010).Grazing can also precipitate species shift from long-lived, slow growing species, to faster growing pioneer species that are better adapted to grazing (Kelkar, Arthur, Marba, & Alcoverro, 2013;Knapp et al., 1999). Additionally, grazing on below-ground plant parts can enhance species diversity by creating regeneration niches through sediment disturbance (Hidding, Nolet, Boer, Vries, & Klaassen, 2010).The impact of herbivory on plant coexistence has been shown across diverse ecosystems, including examples from terrestrial and aquatic systems (Augustine & McNaughton, 1998;Bakker, Pagès, Arthur, & Alcoverro, 2015). This being the case, there is a potential for grazers to increase the success of invasive plants. Evidence of interactions between grazing and invasive plants go both ways.Nonnative species may come to dominate ...

General rightsIt is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an open content license (like Creative Commons). Disclaimer/Complaints regulationsIf you believe that digital publication of certain material infringes any of your rights or (privacy) interests, please let the Library know, stating your reasons. In case of a legitimate complaint, the Library will make the material inaccessible and/or remove it from the website. Please Ask the Library: http://uba.uva.nl/en/contact, or a letter to: Library of the University of Amsterdam, Secretariat, Singel 425, 1012 WP Amsterdam, The Netherlands. You will be contacted as soon as possible. The non-native seagrass species Halophila stipulacea has spread throughout the Eastern Caribbean since 2002, and could potentially impact the functioning of local seagrass ecosystems. Important characteristics for invasiveness, such as dispersal, recruitment and expansion of H. stipulacea at a local scale, are unknown. We assessed H. stipulacea expansion rates within Lac Bay, Bonaire, Dutch Caribbean (7 km 2 ), since its establishment in 2010 and tested the settlement potential of uprooted vegetative fragments of H. stipulacea. Using 49 fixed locations, we observed that between 2011 and 2015 the occurrence of H. stipulacea in the bay increased significantly from 6% to 20% while native Thalassia testudinum occurrence decreased significantly from 53% to 33%. Free-floating H. stipulacea fragments that were collected and tethered above the sediment rooted within 10 days with a settlement success rate of 100%. The growth of settled fragments was on average 0.91 shoots d −1 . The ongoing shift from native T. testudinum to introduced H. stipulacea dominated meadows may have important consequences for multiple Caribbean seagrass ecosystem functions. Given the large difference in size between the two seagrass species, functions such as coastal protection, habitat structure, food availability, and the stability and resilience of these systems can be altered. The next steps towards modelling future expansion of H. stipulacea throughout the Caribbean and beyond should include the assessment of fragment viability and dispersal distance, and the impacts of natural and anthropogenic disturbance on vegetative fragment density, dispersion and settlement by this species. ARTICLE HISTORY

1. Increasing green turtle abundance will lead to increased grazing within seagrass habitats-ecosystems that are important for carbon sequestration and storage.However, it is not well understood how carbon dynamics in these ecosystems respond to grazing and whether a response differs among meadows or locations.2. We measured seagrass ecosystem metabolism in grazed and ungrazed areas of Thalassia testudinum meadows with established green turtle foraging areas across the Greater Caribbean and Gulf of Mexico. We sampled meadows from five locations that differed in seagrass and environmental characteristics. Established meadows of the invasive seagrass Halophila stipulacea were also present at two of these locations, and we measured ecosystem metabolism in these meadows for comparison to grazed and ungrazed areas of the native T. testudinum.3. Across all individual sites, rates of net ecosystem production (NEP) ranged from 56% to 96% lower in grazed areas than ungrazed areas of T. testudinum meadows.Rates of NEP were also strongly, positively correlated with above-ground seagrass biomass across sites. While metabolic carbon capture rates were lower in grazed areas, heterotrophic respiration was not stimulated, and grazing therefore did not result in significant metabolic remineralization of carbon in these meadows. NEP in H. stipulacea meadows was similar to rates in T. testudinum meadows at all three sites, suggesting that metabolic carbon capture may remain similar in Caribbean meadows where this invasive seagrass is replacing native species. Synthesis.Our results show that there is a consistent response in metabolic carbon dynamics to green turtle grazing in T. testudinum meadows across the Greater Caribbean region. An increase in grazing will not likely stimulate remineralization of carbon as these important habitats are returned to a natural grazed state. K E Y W O R D Scarbon dynamics, grazing, green turtle, Halophila stipulacea, invasive, metabolism, plantherbivore interactions, Thalassia testudinum

Understanding how megaherbivores incorporate habitat features into their foraging behavior is key toward understanding how herbivores shape the surrounding landscape. While the role of habitat structure has been studied within the context of predator-prey dynamics and grazing behavior in terrestrial systems, there is a limited understanding of how structure influences megaherbivore grazing in marine ecosystems. To investigate the response of megaherbivores (green turtles) to habitat features, we experimentally introduced structure at two spatial scales in a shallow seagrass meadow in The Bahamas. Turtle density increased 50-fold (to 311 turtles ha À1 ) in response to the structures, and turtles were mainly grazing and resting (low vigilance behavior). This resulted in a grazing patch exceeding the size of the experimental setup (242 m 2 ), with reduced seagrass shoot density and aboveground biomass. After structure removal, turtle density decreased and vigilance increased (more browsing and shorter surfacing times), while seagrass within the patch partly recovered. Even at a small scale (9 m 2 ), artificial structures altered turtle grazing behavior, resulting in grazing patches in 60% of the plots.Our results demonstrate that marine megaherbivores select habitat features as foraging sites, likely to be a predator refuge, resulting in heterogeneity in seagrass bed structure at the landscape scale.

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