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(PDF) Interactive effects of herbivory and substrate orientation on algal community dynamics on a coral reef | Alain Duran - Academia.edu
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We experimentally evaluated" /> <title>(PDF) Interactive effects of herbivory and substrate orientation on algal community dynamics on a coral reef | Alain Duran - Academia.edu</title> <link rel="canonical" href="https://www.academia.edu/37570052/Interactive_effects_of_herbivory_and_substrate_orientation_on_algal_community_dynamics_on_a_coral_reef" /> <script async src="https://www.googletagmanager.com/gtag/js?id=G-5VKX33P2DS"></script> <script> window.dataLayer = window.dataLayer || []; function gtag(){dataLayer.push(arguments);} gtag('js', new Date()); gtag('config', 'G-5VKX33P2DS', { cookie_domain: 'academia.edu', send_page_view: false, }); gtag('event', 'page_view', { 'controller': "single_work", 'action': "show", 'controller_action': 'single_work#show', 'logged_in': 'false', 'edge': 'unknown', // Send nil if there is no A/B test bucket, in case some records get logged // with missing data - that way we can distinguish between the two cases. // ab_test_bucket should be of the form <ab_test_name>:<bucket> 'ab_test_bucket': null, }) </script> <script> var $controller_name = 'single_work'; var $action_name = "show"; var $rails_env = 'production'; var $app_rev = '6c4644673ab7a694ba8b49783a31f1c6511188e3'; var $domain = 'academia.edu'; var $app_host = "academia.edu"; var $asset_host = "academia-assets.com"; var $start_time = new Date().getTime(); var $recaptcha_key = "6LdxlRMTAAAAADnu_zyLhLg0YF9uACwz78shpjJB"; var $recaptcha_invisible_key = "6Lf3KHUUAAAAACggoMpmGJdQDtiyrjVlvGJ6BbAj"; var $disableClientRecordHit = false; </script> <script> window.require = { config: function() { return function() {} } } </script> <script> window.Aedu = window.Aedu || {}; window.Aedu.hit_data = null; window.Aedu.serverRenderTime = new Date(1733363012000); window.Aedu.timeDifference = new Date().getTime() - 1733363012000; </script> <script type="application/ld+json">{"@context":"https://schema.org","@type":"ScholarlyArticle","abstract":"Herbivory is a significant driver of algal community dynamics on coral reefs. However, abiotic factors such as the complexity and orientation of the benthos often mediate the impact of herbivores on benthic communities. We experimentally evaluated the independent and interactive effects of substrate orientation and herbivorous fishes on algal community dynamics on a coral reef in the Florida Keys, USA. We created horizontal and vertical substrates, mimicking the trend in the reduction of vertical surfaces of coral reefs, to assess how algal communities developed either with herbivory (open areas) or without herbivory (herbivore exclosures). We found that substrate orientation was the dominant influence on macroalgal community composition. Herbivores had little impact on community development of vertical substrates as crustose algae dominated these substrates regardless of being in exclosures or open areas. In contrast, herbivores strongly impacted communities on horizontal substrates, with upright macroalgae (e.g., Dictyota spp., articulated coralline algae) dominating herbivore exclo-sures, while filamentous turf algae and sediment dominated open areas. Outside of exclosures, differences between vertical and horizontal substrates exposed to herbivores persisted despite similar intensity of herbivory. Our results suggest that the orientation of the reef benthos has an important impact on benthic communities. On vertical surfaces, abiotic factors may be more important for structuring algal communities while herbivory may be more important for controlling algal dynamics in flatter areas. Thus, the decline in structural complexity of Caribbean coral reefs and the flattening of reef substrates may fundamentally alter the impact that herbivores have on benthic community dynamics.","author":[{"@context":"https://schema.org","@type":"Person","name":"Alain Duran"}],"contributor":[],"dateCreated":"2018-10-11","dateModified":"2018-10-11","headline":"Interactive effects of herbivory and substrate orientation on algal community dynamics on a coral reef","image":"https://attachments.academia-assets.com/57547375/thumbnails/1.jpg","inLanguage":"en","keywords":["Coral Reefs","Herbivory","Structural Complexity","Algal communities"],"publisher":{"@context":"https://schema.org","@type":"Organization","name":null},"sourceOrganization":[{"@context":"https://schema.org","@type":"EducationalOrganization","name":"fiu"}],"thumbnailUrl":"https://attachments.academia-assets.com/57547375/thumbnails/1.jpg","url":"https://www.academia.edu/37570052/Interactive_effects_of_herbivory_and_substrate_orientation_on_algal_community_dynamics_on_a_coral_reef"}</script><link rel="stylesheet" media="all" href="//a.academia-assets.com/assets/single_work_page/loswp-102fa537001ba4d8dcd921ad9bd56c474abc201906ea4843e7e7efe9dfbf561d.css" /><link rel="stylesheet" media="all" href="//a.academia-assets.com/assets/design_system/body-8d679e925718b5e8e4b18e9a4fab37f7eaa99e43386459376559080ac8f2856a.css" /><link rel="stylesheet" media="all" href="//a.academia-assets.com/assets/design_system/button-3cea6e0ad4715ed965c49bfb15dedfc632787b32ff6d8c3a474182b231146ab7.css" /><link rel="stylesheet" media="all" 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[{"id":57547375,"identifier":"Attachment_57547375","shouldShowBulkDownload":false}]; window.loswp.shouldDetectTimezone = true; window.loswp.shouldShowBulkDownload = true; window.loswp.showSignupCaptcha = false window.loswp.willEdgeCache = false; window.loswp.work = {"work":{"id":37570052,"created_at":"2018-10-11T13:21:00.068-07:00","from_world_paper_id":null,"updated_at":"2021-01-16T12:40:36.394-08:00","_data":{"abstract":"Herbivory is a significant driver of algal community dynamics on coral reefs. However, abiotic factors such as the complexity and orientation of the benthos often mediate the impact of herbivores on benthic communities. We experimentally evaluated the independent and interactive effects of substrate orientation and herbivorous fishes on algal community dynamics on a coral reef in the Florida Keys, USA. We created horizontal and vertical substrates, mimicking the trend in the reduction of vertical surfaces of coral reefs, to assess how algal communities developed either with herbivory (open areas) or without herbivory (herbivore exclosures). We found that substrate orientation was the dominant influence on macroalgal community composition. Herbivores had little impact on community development of vertical substrates as crustose algae dominated these substrates regardless of being in exclosures or open areas. In contrast, herbivores strongly impacted communities on horizontal substrates, with upright macroalgae (e.g., Dictyota spp., articulated coralline algae) dominating herbivore exclo-sures, while filamentous turf algae and sediment dominated open areas. Outside of exclosures, differences between vertical and horizontal substrates exposed to herbivores persisted despite similar intensity of herbivory. Our results suggest that the orientation of the reef benthos has an important impact on benthic communities. On vertical surfaces, abiotic factors may be more important for structuring algal communities while herbivory may be more important for controlling algal dynamics in flatter areas. Thus, the decline in structural complexity of Caribbean coral reefs and the flattening of reef substrates may fundamentally alter the impact that herbivores have on benthic community dynamics."},"document_type":"paper","pre_hit_view_count_baseline":null,"quality":"high","language":"en","title":"Interactive effects of herbivory and substrate orientation on algal community dynamics on a coral reef","broadcastable":true,"draft":false,"has_indexable_attachment":true,"indexable":true}}["work"]; window.loswp.workCoauthors = [47292361]; window.loswp.locale = "en"; window.loswp.countryCode = "SG"; window.loswp.cwvAbTestBucket = ""; window.loswp.designVariant = "ds_vanilla"; window.loswp.fullPageMobileSutdModalVariant = "control"; window.loswp.useOptimizedScribd4genScript = false; window.loswp.appleClientId = 'edu.academia.applesignon';</script><script defer="" src="https://accounts.google.com/gsi/client"></script><div class="ds-loswp-container"><div class="ds-work-card--grid-container"><div class="ds-work-card--container js-loswp-work-card"><div class="ds-work-card--cover"><div class="ds-work-cover--wrapper"><div class="ds-work-cover--container"><button class="ds-work-cover--clickable js-swp-download-button" data-signup-modal="{"location":"swp-splash-paper-cover","attachmentId":57547375,"attachmentType":"pdf"}"><img alt="First page of “Interactive effects of herbivory and substrate orientation on algal community dynamics on a coral reef”" class="ds-work-cover--cover-thumbnail" src="https://0.academia-photos.com/attachment_thumbnails/57547375/mini_magick20190110-31880-zsry6h.png?1547169502" /><img alt="PDF Icon" class="ds-work-cover--file-icon" src="//a.academia-assets.com/assets/single_work_splash/adobe.icon-574afd46eb6b03a77a153a647fb47e30546f9215c0ee6a25df597a779717f9ef.svg" /><div class="ds-work-cover--hover-container"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">download</span><p>Download Free PDF</p></div><div class="ds-work-cover--ribbon-container">Download Free PDF</div><div class="ds-work-cover--ribbon-triangle"></div></button></div></div></div><div class="ds-work-card--work-information"><h1 class="ds-work-card--work-title">Interactive effects of herbivory and substrate orientation on algal community dynamics on a coral reef</h1><div class="ds-work-card--work-authors ds-work-card--detail"><a class="ds-work-card--author js-wsj-grid-card-author ds2-5-body-md ds2-5-body-link" data-author-id="47292361" href="https://fiu.academia.edu/ADuran"><img alt="Profile image of Alain Duran" class="ds-work-card--author-avatar" src="https://0.academia-photos.com/47292361/14742967/19500547/s65_alain.duran.jpg" />Alain Duran</a></div><div class="ds-work-card--detail"></div><p class="ds-work-card--work-abstract ds-work-card--detail ds2-5-body-md">Herbivory is a significant driver of algal community dynamics on coral reefs. However, abiotic factors such as the complexity and orientation of the benthos often mediate the impact of herbivores on benthic communities. We experimentally evaluated the independent and interactive effects of substrate orientation and herbivorous fishes on algal community dynamics on a coral reef in the Florida Keys, USA. We created horizontal and vertical substrates, mimicking the trend in the reduction of vertical surfaces of coral reefs, to assess how algal communities developed either with herbivory (open areas) or without herbivory (herbivore exclosures). We found that substrate orientation was the dominant influence on macroalgal community composition. Herbivores had little impact on community development of vertical substrates as crustose algae dominated these substrates regardless of being in exclosures or open areas. In contrast, herbivores strongly impacted communities on horizontal substrates, with upright macroalgae (e.g., Dictyota spp., articulated coralline algae) dominating herbivore exclo-sures, while filamentous turf algae and sediment dominated open areas. Outside of exclosures, differences between vertical and horizontal substrates exposed to herbivores persisted despite similar intensity of herbivory. Our results suggest that the orientation of the reef benthos has an important impact on benthic communities. On vertical surfaces, abiotic factors may be more important for structuring algal communities while herbivory may be more important for controlling algal dynamics in flatter areas. Thus, the decline in structural complexity of Caribbean coral reefs and the flattening of reef substrates may fundamentally alter the impact that herbivores have on benthic community dynamics.</p><div class="ds-work-card--button-container"><button class="ds2-5-button js-swp-download-button" data-signup-modal="{"location":"continue-reading-button--work-card","attachmentId":57547375,"attachmentType":"pdf","workUrl":"https://www.academia.edu/37570052/Interactive_effects_of_herbivory_and_substrate_orientation_on_algal_community_dynamics_on_a_coral_reef"}">See full PDF</button><button class="ds2-5-button ds2-5-button--secondary js-swp-download-button" data-signup-modal="{"location":"download-pdf-button--work-card","attachmentId":57547375,"attachmentType":"pdf","workUrl":"https://www.academia.edu/37570052/Interactive_effects_of_herbivory_and_substrate_orientation_on_algal_community_dynamics_on_a_coral_reef"}"><span class="material-symbols-outlined" style="font-size: 20px" 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free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/893495/Nutrient_versus_herbivore_control_of_macroalgal_community_development_and_coral_growth_on_a_Caribbean_reef"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="5" data-entity-id="31882781" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/31882781/Coral_reef_grazer_benthos_dynamics_complicated_by_invasive_algae_in_a_small_marine_reserve_OPEN">Coral reef grazer-benthos dynamics complicated by invasive algae in a small marine reserve OPEN</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="27371925" href="https://hawaii.academia.edu/AlanFriedlander">Alan Friedlander</a><span>, </span><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32909706" href="https://curtin.academia.edu/KostantinosStamoulis">Kostantinos A Stamoulis</a></div><p class="ds-related-work--abstract ds2-5-body-sm">Blooms of alien invasive marine algae have become common, greatly altering the health and stability of nearshore marine ecosystems. Concurrently, herbivorous fishes have been severely overfished in many locations worldwide, contributing to increases in macroalgal cover. We used a multi-pronged, interdisciplinary approach to test if higher biomass of herbivorous fishes inside a no-take marine reserve makes this area more resistant to invasive algal overgrowth. Over a two year time period, we (1) compared fish biomass and algal cover between two fished and one unfished patch reef in Hawai'i, (2) used acoustic telemetry to determine fidelity of herbivorous fishes to the unfished reef, and (3) used metabarcoding and next-generation sequencing to determine diet composition of herbivorous fishes. Herbivore fish biomass was significantly higher in the marine reserve compared to adjacent fished reefs, whereas invasive algal cover differed by species. Herbivorous fish movements were largely confined to the unfished patch reef where they were captured. Diet analysis indicated that the consumption of invasive algae varied among fish species, with a high prevalence of comparatively rare native algal species. Together these findings demonstrate that the contribution of herbivores to coral reef resilience, via resistance to invasive algae invasion, is complex and species-specific. Herbivorous fishes and sea urchins are primarily responsible for the high grazing intensity found in healthy coral reef ecosystems 1–3. This intense grazing pressure has a large influence on the distribution of algae on coral reefs, with macroalgae generally rare in reef zones with the highest grazing pressure 2,4,5. Although there has been some debate regarding the mechanisms and causality of competition between algae and corals 6 high algal biomass has been shown to have a negative effect on coral health 7–9. Based on their role in algal removal, herbivorous fishes are considered to promote reef resilience and assist in reef recovery to coral dominated states after a disturbance 10–12. Over the past several decades, blooms of both indigenous and introduced marine algae have become common on coral reefs, greatly altering the health and stability of nearshore ecosystems 10,13. On reefs subjected to anthropogenic disturbances such as increased terrestrial nutrient inputs or the removal of grazers by overfishing, algal growth rates may exceed grazing rates, resulting in overgrowth of hard corals and other non-mobile ben-thic invertebrates, and suppression of coral recruitment 14–16. Herbivorous fishes, particularly large parrotfishes and surgeonfishes, have been severely overfished in many locations/regions and reduction in these grazers has been thought to contribute to increases in macroalgae and subsequent decreases in coral cover 17–19. Introduced species of macroalgae are often not subject to ecological controls that normally limit abundance in their native range, such as high grazing pressure from native herbivores, allowing them to become invasive 20,21 and accelerate coral-algal phase shifts on coral reefs. Herbivorous fishes play an important role in promoting resilience and supporting coral recovery, and tend to have small home ranges, suggesting that relatively small-scale (1–10 s km) variation in their abundance will contribute to local changes in rates of reef recovery 22,23. Therefore, local-scale management measures that decrease fishing mortality and increase the abundance and size of herbivorous fishes, such as marine reserves, are expected to play a significant role in supporting recovery and resilience of coral reef ecosystems 24,25. The extent to which different species of herbivorous fishes control macroalgae and promote coral recovery will also depend on their functional role and the algae upon which they graze 26. Herbivorous fish species tend to be selective in their</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Coral reef grazer-benthos dynamics complicated by invasive algae in a small marine reserve OPEN","attachmentId":52168773,"attachmentType":"pdf","work_url":"https://www.academia.edu/31882781/Coral_reef_grazer_benthos_dynamics_complicated_by_invasive_algae_in_a_small_marine_reserve_OPEN","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/31882781/Coral_reef_grazer_benthos_dynamics_complicated_by_invasive_algae_in_a_small_marine_reserve_OPEN"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="6" data-entity-id="11830233" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/11830233/Herbivores_strongly_influence_algal_recruitment_in_both_coral_and_algal_dominated_coral_reef_habitats">Herbivores strongly influence algal recruitment in both coral-and algal-dominated coral reef habitats</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="29187443" href="https://degois.academia.edu/DavidAbecasis">David Abecasis</a></div><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" 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class="ds-related-work--container js-wsj-grid-card" data-collection-position="7" data-entity-id="47317398" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/47317398/Herbivory_Algal_Distribution_and_the_Maintenance_of_Between_Habitat_Diversity_on_a_Tropical_Fringing_Reef">Herbivory, Algal Distribution, and the Maintenance of Between-Habitat Diversity on a Tropical Fringing Reef</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32057315" href="https://gatech.academia.edu/MarkHay">Mark Hay</a></div><p class="ds-related-work--metadata ds2-5-body-xs">The American Naturalist, 1981</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Herbivory, Algal Distribution, and the Maintenance of Between-Habitat Diversity on a Tropical Fringing Reef","attachmentId":66473464,"attachmentType":"pdf","work_url":"https://www.academia.edu/47317398/Herbivory_Algal_Distribution_and_the_Maintenance_of_Between_Habitat_Diversity_on_a_Tropical_Fringing_Reef","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/47317398/Herbivory_Algal_Distribution_and_the_Maintenance_of_Between_Habitat_Diversity_on_a_Tropical_Fringing_Reef"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="8" data-entity-id="13102076" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/13102076/Phase_shifts_and_the_role_of_herbivory_in_the_resilience_of_coral_reefs">Phase shifts and the role of herbivory in the resilience of coral reefs</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32352850" href="https://newcastle.academia.edu/NicholasPolunin">Nicholas Polunin</a><span>, </span><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32422544" href="https://newcastle.academia.edu/JBythell">J. Bythell</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Coral Reefs, 2007</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Phase shifts and the role of herbivory in the resilience of coral reefs","attachmentId":45665285,"attachmentType":"pdf","work_url":"https://www.academia.edu/13102076/Phase_shifts_and_the_role_of_herbivory_in_the_resilience_of_coral_reefs","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/13102076/Phase_shifts_and_the_role_of_herbivory_in_the_resilience_of_coral_reefs"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="9" data-entity-id="78405828" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/78405828/The_benefits_of_herbivory_outweigh_the_costs_of_bioerosion_in_a_eutrophic_coral_community">The benefits of herbivory outweigh the costs of bioerosion in a eutrophic coral community</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="26364968" href="https://hku-hk.academia.edu/BaydenRussell">Bayden D Russell</a></div><p class="ds-related-work--metadata ds2-5-body-xs">2021</p><p class="ds-related-work--abstract ds2-5-body-sm">Herbivores play an integral part in maintaining the health of coral reefs by suppressing the growth of algae and accumulation of sediment and facilitating coral growth. However, in predator-depleted systems where densities of herbivores are unnaturally high, grazing can have detrimental effects on corals through excessive bioerosion. Yet, these benefits and costs are rarely investigated concurrently, especially in eutrophic systems where grazers may play a disproportionate role. We used a year-long exclusion experiment to elucidate the effect of natural densities of the dominant herbivore (the sea urchin Diadema setosum) on coral communities in a heavily fished and eutrophic system (Hong Kong, China). To assess benthic community response to grazing, we monitored the survival and growth of three locally abundant coral species (Pavona decussata, Platygyra carnosus and Porites sp.), algal and sediment accumulation, and bioerosion of coral skeletons across seasons. We found that urchins...</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"The benefits of herbivory outweigh the costs of bioerosion in a eutrophic coral community","attachmentId":85464862,"attachmentType":"pdf","work_url":"https://www.academia.edu/78405828/The_benefits_of_herbivory_outweigh_the_costs_of_bioerosion_in_a_eutrophic_coral_community","alternativeTracking":true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/78405828/The_benefits_of_herbivory_outweigh_the_costs_of_bioerosion_in_a_eutrophic_coral_community"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div></div></div><div class="ds-sticky-ctas--wrapper js-loswp-sticky-ctas hidden"><div class="ds-sticky-ctas--grid-container"><div class="ds-sticky-ctas--container"><button class="ds2-5-button js-swp-download-button" data-signup-modal="{"location":"continue-reading-button--sticky-ctas","attachmentId":57547375,"attachmentType":"pdf","workUrl":null}">See full PDF</button><button class="ds2-5-button ds2-5-button--secondary js-swp-download-button" data-signup-modal="{"location":"download-pdf-button--sticky-ctas","attachmentId":57547375,"attachmentType":"pdf","workUrl":null}"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">download</span>Download PDF</button></div></div></div><div class="ds-below-fold--grid-container"><div class="ds-work--container js-loswp-embedded-document"><div class="attachment_preview" data-attachment="Attachment_57547375" style="display: none"><div class="js-scribd-document-container"><div class="scribd--document-loading js-scribd-document-loader" style="display: block;"><img alt="Loading..." src="//a.academia-assets.com/images/loaders/paper-load.gif" /><p>Loading Preview</p></div></div><div style="text-align: center;"><div class="scribd--no-preview-alert js-preview-unavailable"><p>Sorry, preview is currently unavailable. 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Proceedings of the 10th International Coral Reef Symposium, Okinawa 3:854-861</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="46542" href="https://uvi.academia.edu/DrTylerSmith">Tyler Smith</a></div><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{"location":"wsj-grid-card-download-pdf-modal","work_title":"Smith T (2006) Benthic algal production in an upwelling coral reef environment (Gulf of Panamá): Evidence for control by fish herbivores. 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