Mesophotic corals in Hawai'i maintain autotrophy to survive low-light conditions.

In mesophotic coral ecosystems, reef-building corals and their photosynthetic symbionts can survive with less than 1% of surface irradiance. How depth-specialist corals rely upon autotrophically and heterotrophically derived energy sources across the mesophotic zone remains unclear. We analysed the stable carbon (δ13C) and nitrogen (δ15N) isotope values of a Leptoseris community from the 'Au'au Channel, Maui, Hawai'i (65-125 m) including four coral host species living symbiotically with three algal haplotypes. We characterized the isotope values of hosts and symbionts across species and depth to compare trophic strategies. Symbiont δ13C was consistently 0.5‰ higher than host δ13C at all depths. Mean colony host and symbiont δ15N differed by up to 3.7‰ at shallow depths and converged at deeper depths. These results suggest that both heterotrophy and autotrophy remained integral to colony survival across depth. The increasing similarity between host and symbiont δ15N at deeper depths suggests that nitrogen is more efficiently shared between mesophotic coral hosts and their algal symbionts to sustain autotrophy. Isotopic trends across depth did not generally vary by host species or algal haplotype, suggesting that photosynthesis remains essential to Leptoseris survival and growth despite low light availability in the mesophotic zone.

Organellar genomic characterization of Anunuuluaehu liula representing a new genus and species of Phyllophoraceae (Gigartinales, Rhodophyta) from the mesophotic zone of Hawai'i.

Over the last 2 decades, routine collections in the Hawaiian Archipelago have expanded to mesophotic reefs, leading to the discovery of a new red algal genus and species, here described as Anunuuluaehu liula gen. et sp. nov. This study provides a detailed genus and species description and characterizes chloroplast and mitochondrial organellar genomes. The new genus, Anunuuluaehu, shares many characteristics with the family Phyllophoraceae and shows close similarities to Archestennogramma and Stenogramma, including habit morphology, nemathecia forming proliferations at the outer cortex with terminal chains of tetrasporangia, and carposporophytes with multi-layered pericarps. The single species in this genus exhibits distinctive features within the Phyllophoraceae: the presence of single-layer construction of large medullary cells and the development of long, tubular gonimoblastic filaments. Multi-gene phylogenetic analyses confirmed it as a unique, monophyletic lineage within the family. Cis-splicing genes, interrupted by intron-encoded proteins within group II introns, are present in both the chloroplast and mitochondrial genomes of A. liula. Notably, a specific region of the coxI group II intron exhibits similarity to fungal introns. Anunuuluaehu liula is presumed to be endemic to the Hawaiian Archipelago and thus far is known to live solely at mesophotic depths from Holaniku to Kaho'olawe ranging from 54 to 201 m, which is the deepest collection record of any representative in the family. Overall, this study enhances our understanding of the genomic and taxonomic complexities of red algae in mesophotic habitats, emphasizing the significance of continued research in this area to uncover further insights into evolutionary processes and biogeographic patterns.

Divergent responses of native and invasive macroalgae to submarine groundwater discharge.

Marine macroalgae are important indicators of healthy nearshore groundwater dependent ecosystems (GDEs), which are emergent global conservation priorities. Submarine groundwater discharge (SGD) supports abundant native algal communities in GDEs via elevated but naturally derived nutrients. GDEs are threatened by anthropogenic nutrient inputs that pollute SGD above ambient levels, favoring invasive algae. Accordingly, this case study draws on the GDE conditions of Kona, Hawai'i where we evaluated daily photosynthetic production and growth for two macroalgae; a culturally valued native (Ulva lactuca) and an invasive (Hypnea musciformis). Manipulative experiments-devised to address future land-use, climate change, and water-use scenarios for Kona-tested algal responses under a natural range of SGD nutrient and salinity levels. Our analyses demonstrate that photosynthesis and growth in U. lactuca are optimal in low-salinity, high-nutrient waters, whereas productivity for H. musciformis appears limited to higher salinities despite elevated nutrient subsidies. These findings suggest that reductions in SGD via climate change decreases in rainfall or increased water-use from the aquifer may relax physiological constraints on H. musciformis. Collectively, this study reveals divergent physiologies of a native and an invasive macroalga to SGD and highlights the importance of maintaining SGD quantity and quality to protect nearshore GDEs.

Protocol to control the invasive alga Avrainvillea lacerata in a shallow Hawaiian reef flat.

Premise: A novel control technique was developed to mitigate an invasive siphonous green alga, Avrainvillea lacerata (Dichotomosiphonaceae), within a shallow degraded reef flat in O'ahu, Hawai'i. Methods and Results: Replicated treatments of 3% and 10% hydrogen peroxide (H2O2) were administered into individual basal attachments of the bed-forming invasive seaweed on the Paiko reef, O'ahu. Relative electron transport rate maxima (rETRm) were measured using a Walz Diving Pulse Amplitude Modulated Fluorometer in two replicate 100-m2 plots in 2020. Over the period of this short-term study, rETRm decreased following injections of either concentration of H2O2 in contrast with negative and positive controls. Conclusions: Compared with existing techniques that have used oxidizing agents in the marine environment in localized areas, the protocol described here has the potential to successfully decrease macroalgal carbon gain, potentially leading to loss of biomass at larger scales.

Introduced mangroves escape damage from marine and terrestrial enemies.

The enemy release hypothesis (ERH) posits that introduced species often leave their enemies behind when introduced to a new range. This release from enemies may allow introduced species to achieve higher growth and reproduction and may explain why some invaders flourish in new locations. Red mangroves (Rhizophora mangle) were introduced to Hawai'i from Florida over a century ago. Because Hawai'i has no native mangroves, the arrival of R. mangle fundamentally changed the structure and function of estuarine shorelines. While numerous enemies affect red mangroves in their native range (tropical America), in Hawai'i, mangroves apparently experience little herbivory, which may explain why introduced mangroves are so productive, fecund, and continue to spread. In this study, we compared the effects of enemies in native and introduced populations of brackish red mangroves (R. mangle) in 8-10 sites in the native range (Florida, Belize, and Panama) and introduced range of mangroves (Hawai'i). At each site, we measured the (1) occurrence of enemies using timed visual surveys, (2) occurrence of damage to different mangrove structures (leaves, apical buds, dead twigs, roots, propagules, and seedlings), and (3) rate of propagule herbivory using tethering experiments. Consistent with the ERH, we found an order of magnitude less damage and fewer enemies in introduced than native mangrove sites. While introduced mangroves harbored few enemies and minimal damage, native mangroves were affected by numerous enemies, including leaf-eating crabs, specialist bud moths, wood-boring insects and isopods, and propagule predators. These patterns were consistent across all plant structures (roots to leaves), among marine and terrestrial enemies, and across functional groups (browsers, borers, pathogens, etc.), which demonstrates enemy escape occurs consistently among different functional groups and via trophic (e.g., herbivores) and non-trophic (e.g., root borers) interactions. Our study is among the first biogeographical enemy release studies to take a comprehensive approach to quantifying the occurrence of damage from a broad suite of marine and terrestrial taxa across an array of wetland plant structures. Understanding how natural enemies alter this key foundation species will become increasingly relevant globally as mangroves continue to invade new regions through intentional plantings or range expansion driven by climate change.

Taxonomic determination of the cryptogenic red alga, Chondria tumulosa sp. nov., (Rhodomelaceae, Rhodophyta) from Papahanaumokuakea Marine National Monument, Hawai'i, USA: A new species displaying invasive characteristics.

Survey cruises by the National Oceanic and Atmospheric Administration (NOAA) in 2016 and 2019 yielded specimens of an undetermined red alga that rapidly attained alarming levels of benthic coverage at Pearl and Hermes Atoll, Papahanaumokuakea Marine National Monument, Hawai'i. By 2019 the seaweed had covered large expanses on the northeast side of the atoll with mat-like, extensive growth of entangled thalli. Specimens were analyzed using light microscopy and molecular analysis, and were compared to morphological descriptions in the literature for closely related taxa. Light microscopy demonstrated that the specimens likely belonged to the rhodomelacean genus Chondria, yet comparisons to taxonomic literature revealed no morphological match. DNA sequence analyses of the mitochondrial COI barcode marker, the plastidial rbcL gene, and the nuclear SSU gene confirmed its genus-level placement and demonstrated that this alga was unique compared to all other available sequences. Based on these data, this cryptogenic seaweed is here proposed as a new species: Chondria tumulosa A.R.Sherwood & J.M.Huisman sp. nov. Chondria tumulosa is distinct from all other species of Chondria based on its large, robust thalli, a mat-forming tendency, large axial diameter in mature branches (which decreases in diameter with subsequent orders of branching), terete axes, and bluntly rounded apices. Although C. tumulosa does not meet the criteria for the definition of an invasive species given that it has not been confirmed as introduced to Pearl and Hermes Atoll, this seaweed is not closely related to any known Hawaiian native species and is of particular concern given its sudden appearance and rapid increase in abundance in the Papahanaumokuakea Marine National Monument; an uninhabited, remote, and pristine island chain to the northwest of the Main Hawaiian Islands.

Herbaria macroalgae as a proxy for historical upwelling trends in Central California.

Planning for future ocean conditions requires historical data to establish more informed ecological baselines. To date, this process has been largely limited to instrument records and observations that begin around 1950. Here, we show how marine macroalgae specimens from herbaria repositories may document long-term ecosystem processes and extend historical information records into the nineteenth century. We tested the effect of drying and pressing six macroalgae species on amino acid, heavy metal and bulk stable isotope values over 1 year using modern and archived paper. We found historical paper composition did not consistently affect values. Certain species, however, had higher variability in particular metrics while others were more consistent. Multiple herbaria provided Gelidium (Rhodophyta) samples collected in southern Monterey Bay from 1878 to 2018. We examined environmental relationships and found δ15N correlated with the Bakun upwelling index, the productivity regime of this ecosystem, from 1946 to 2018. Then, we hindcasted the Bakun index using its derived relationship with Gelidium δ15N from 1878 to 1945. This hindcast provided new information, observing an upwelling decrease mid-century leading up to the well-known sardine fishery crash. Our case study suggests marine macroalgae from herbaria are an underused resource of the marine environment that precedes modern scientific data streams.

Algal bioassays detect modeled loading of wastewater-derived nitrogen in coastal waters of O'AHU, HAWAI'I.

Previous studies indicate coastlines are at risk of wastewater contamination from injection wells, cesspools, and septic systems. In this study, common marine algae were used to ground-truth modeled loading of wastewater-derived N to coastlines of O'ahu, Hawai'i. Macroalgae were collected and/or deployed at 118 sites and analyzed for tissue δ15N and N %. Wastewater source locations were used to estimate wastewater-derived N in groundwater with the modeling software MT3DMS/MODFLOW. Algal bioassays identified six coastal regions subjected to elevated wastewater-derived N loading. In a case study, submarine groundwater discharge (estimated by 222Rn mass balance) was related to wastewater loading from onsite sewage disposal systems (OSDS) and municipal wastewater injection wells in Waimanalo. The highest 222Rn-derived SGD rate and N flux were 21.4 m3/m/d and 62.6 g/m/d, respectively. The results of this study suggest that OSDS and injection wells discharge substantial volumes of wastewater and N across broad regions of coastal O'ahu.

Recognizing and Reducing Barriers to Science and Math Education and STEM Careers for Native Hawaiians and Pacific Islanders.

Climate change is impacting the Pacific Islands first and most drastically, yet few native islanders are trained to recognize, analyze, or mitigate the impacts in these islands. To understand the reasons why low numbers of Native Hawaiians and Pacific Islanders enter colleges, enroll in science, technology, engineering, and mathematics (STEM) courses, or undertake life sciences/STEM careers, 25 representatives from colleges and schools in seven U.S.-affiliated states and countries across the Pacific participated in a 2-day workshop. Fourteen were indigenous peoples of their islands. Participants revealed that: 1) cultural barriers, including strong family obligations and traditional and/or religious restrictions, work against students leaving home or entering STEM careers; 2) geographic barriers confront isolated small island communities without secondary schools, requiring students to relocate to a distant island for high school; 3) in many areas, teachers are undertrained in STEM, school science facilities are lacking, and most island colleges lack STEM majors and modern labs; and 4) financial barriers arise, because many islanders must relocate from their home islands to attend high school and college, especially, the costs for moving to Guam, Hawai'i, or the U.S. mainland. Most solutions depend on financial input, but mechanisms to increase awareness of the value of STEM training are also important.

Fungi associated with mesophotic macroalgae from the 'Au'au Channel, west Maui are differentiated by host and overlap terrestrial communities.

Mesophotic coral ecosystems are an almost entirely unexplored and undocumented environment that likely contains vast reservoirs of undescribed biodiversity. Twenty-four macroalgae samples, representing four genera, were collected from a Hawaiian mesophotic reef at water depths between 65 and 86 m in the 'Au'au Channel, Maui, Hawai'i. Algal tissues were surveyed for the presence and diversity of fungi by sequencing the ITS1 gene using Illumina technology. Fungi from these algae were then compared to previous fungal surveys conducted in Hawaiian terrestrial ecosystems. Twenty-seven percent of the OTUs present on the mesophotic coral ecosystem samples were shared between the marine and terrestrial environment. Subsequent analyses indicated that host species of algae significantly differentiate fungal community composition. This work demonstrates yet another understudied habitat with a moderate diversity of fungi that should be considered when estimating global fungal diversity.

Impact of Submarine Groundwater Discharge on Marine Water Quality and Reef Biota of Maui.

Generally unseen and infrequently measured, submarine groundwater discharge (SGD) can transport potentially large loads of nutrients and other land-based contaminants to coastal ecosystems. To examine this linkage we employed algal bioassays, benthic community analysis, and geochemical methods to examine water quality and community parameters of nearshore reefs adjacent to a variety of potential, land-based nutrient sources on Maui. Three common reef algae, Acanthophora spicifera, Hypnea musciformis, and Ulva spp. were collected and/or deployed at six locations with SGD. Algal tissue nitrogen (N) parameters (δ15N, N %, and C:N) were compared with nutrient and δ15N-nitrate values of coastal groundwater and nearshore surface water at all locations. Benthic community composition was estimated for ten 10-m transects per location. Reefs adjacent to sugarcane farms had the greatest abundance of macroalgae, low species diversity, and the highest concentrations of N in algal tissues, coastal groundwater, and marine surface waters compared to locations with low anthropogenic impact. Based on δ15N values of algal tissues, we estimate ca. 0.31 km2 of Kahului Bay is impacted by effluent injected underground at the Kahului Wastewater Reclamation Facility (WRF); this region is barren of corals and almost entirely dominated by colonial zoanthids. Significant correlations among parameters of algal tissue N with adjacent surface and coastal groundwater N indicate that these bioassays provided a useful measure of nutrient source and loading. A conceptual model that uses Ulva spp. tissue δ15N and N % to identify potential N source(s) and relative N loading is proposed for Hawai'i. These results indicate that SGD can be a significant transport pathway for land-based nutrients with important biogeochemical and ecological implications in tropical, oceanic islands.

A comprehensive investigation of mesophotic coral ecosystems in the Hawaiian Archipelago.

Although the existence of coral-reef habitats at depths to 165 m in tropical regions has been known for decades, the richness, diversity, and ecological importance of mesophotic coral ecosystems (MCEs) has only recently become widely acknowledged. During an interdisciplinary effort spanning more than two decades, we characterized the most expansive MCEs ever recorded, with vast macroalgal communities and areas of 100% coral cover between depths of 50-90 m extending for tens of km2 in the Hawaiian Archipelago. We used a variety of sensors and techniques to establish geophysical characteristics. Biodiversity patterns were established from visual and video observations and collected specimens obtained from submersible, remotely operated vehicles and mixed-gas SCUBA and rebreather dives. Population dynamics based on age, growth and fecundity estimates of selected fish species were obtained from laser-videogrammetry, specimens, and otolith preparations. Trophic dynamics were determined using carbon and nitrogen stable isotopic analyses on more than 750 reef fishes. MCEs are associated with clear water and suitable substrate. In comparison to shallow reefs in the Hawaiian Archipelago, inhabitants of MCEs have lower total diversity, harbor new and unique species, and have higher rates of endemism in fishes. Fish species present in shallow and mesophotic depths have similar population and trophic (except benthic invertivores) structures and high genetic connectivity with lower fecundity at mesophotic depths. MCEs in Hawai'i are widespread but associated with specific geophysical characteristics. High genetic, ecological and trophic connectivity establish the potential for MCEs to serve as refugia for some species, but our results question the premise that MCEs are more resilient than shallow reefs. We found that endemism within MCEs increases with depth, and our results do not support suggestions of a global faunal break at 60 m. Our findings enhance the scientific foundations for conservation and management of MCEs, and provide a template for future interdisciplinary research on MCEs worldwide.

New Ulvaceae (Ulvophyceae, Chlorophyta) from mesophotic ecosystems across the Hawaiian Archipelago.

Ulvalean algae (Chlorophyta) are most commonly described from intertidal and shallow subtidal marine environments worldwide, but are less well known from mesophotic environments. Their morphological simplicity and phenotypic plasticity make accurate species determinations difficult, even at the generic level. Here, we describe the mesophotic Ulvales species composition from 13 locations across 2,300 km of the Hawaiian Archipelago. Twenty-eight representative Ulvales specimens from 64 to 125 m depths were collected using technical diving, submersibles, and remotely operated vehicles. Morphological and molecular characters suggest that mesophotic Ulvales in Hawaiian waters form unique communities comprising four species within the genera Ulva and Umbraulva, each with discrete geographic and/or depth-related distributional patterns. Three genetically distinct taxa are supported by both plastid (rbcL and tufA) and nuclear (ITS1) markers, and are presented here as new species: Umbraulva kaloakulau, Ulva ohiohilulu, and Ulva iliohaha. We also propose a new Umbraulva species (Umbraulva kuaweuweu), which is closely related to subtidal records from New Zealand and Australia, but not formally described. To our knowledge, these are the first marine species descriptions from Hawai'i resulting from the collaboration of traditional Hawaiian nomenclature specialists, cultural practitioners and scientists. The difficulty of finding reliable diagnostic morphological characters for these species reflects a common problem worldwide of achieving accurate identification of ulvalean taxa using solely morphological criteria. Mesophotic Ulvales appear to be distinct from shallow-water populations in Hawai'i, but their degree of similarity to mesophotic floras in other locations in the Pacific remains unknown.

Eutrophication and the dietary promotion of sea turtle tumors.

The tumor-forming disease fibropapillomatosis (FP) has afflicted sea turtle populations for decades with no clear cause. A lineage of α-herpesviruses associated with these tumors has existed for millennia, suggesting environmental factors are responsible for its recent epidemiology. In previous work, we described how herpesviruses could cause FP tumors through a metabolic influx of arginine. We demonstrated the disease prevails in chronically eutrophied coastal waters, and that turtles foraging in these sites might consume arginine-enriched macroalgae. Here, we test the idea using High-Performance Liquid Chromatography (HPLC) to describe the amino acid profiles of green turtle (Chelonia mydas) tumors and five common forage species of macroalgae from a range of eutrophic states. Tumors were notably elevated in glycine, proline, alanine, arginine, and serine and depleted in lysine when compared to baseline samples. All macroalgae from eutrophic locations had elevated arginine, and all species preferentially stored environmental nitrogen as arginine even at oligotrophic sites. From these results, we estimate adult turtles foraging at eutrophied sites increase their arginine intake 17-26 g daily, up to 14 times the background level. Arginine nitrogen increased with total macroalgae nitrogen and watershed nitrogen, and the invasive rhodophyte Hypnea musciformis significantly outperformed all other species in this respect. Our results confirm that eutrophication substantially increases the arginine content of macroalgae, which may metabolically promote latent herpesviruses and cause FP tumors in green turtles.

Between tide and wave marks: a unifying model of physical zonation on littoral shores.

The effects of tides on littoral marine habitats are so ubiquitous that shorelines are commonly described as 'intertidal', whereas waves are considered a secondary factor that simply modifies the intertidal habitat. However mean significant wave height exceeds tidal range at many locations worldwide. Here we construct a simple sinusoidal model of coastal water level based on both tidal range and wave height. From the patterns of emergence and submergence predicted by the model, we derive four vertical shoreline benchmarks which bracket up to three novel, spatially distinct, and physically defined zones. The (1) emergent tidal zone is characterized by tidally driven emergence in air; the (2) wave zone is characterized by constant (not periodic) wave wash; and the (3) submergent tidal zone is characterized by tidally driven submergence. The decoupling of tidally driven emergence and submergence made possible by wave action is a critical prediction of the model. On wave-dominated shores (wave height ≫ tidal range), all three zones are predicted to exist separately, but on tide-dominated shores (tidal range ≫ wave height) the wave zone is absent and the emergent and submergent tidal zones overlap substantially, forming the traditional "intertidal zone". We conclude by incorporating time and space in the model to illustrate variability in the physical conditions and zonation on littoral shores. The wave:tide physical zonation model is a unifying framework that can facilitate our understanding of physical conditions on littoral shores whether tropical or temperate, marine or lentic.

Can stormwater be detected by algae in an urban reef in Hawai'i?

Nitrogen (N) enrichment of tropical reefs can result in the dominance of invasive algae. The invasive alga Acanthophora spicifera and the native alga Laurencia nidifica are part of a diverse reef assemblage in 'Ewa Beach, O'ahu. Their N contents and δ(15)N values were investigated to determine if N was enriched and to evaluate potential nitrogenous sources near and removed from storm-drain outlets. δ(15)N values of algae (3.8-17.7‰) were within and above the range for algae around the island (1.9-11.9‰). Elevated algae N isotope values (δ(15)N>+7‰, [N]>1.6%) and seawater nitrate+nitrite levels (0.59-7.93 µM) indicated a mixed, high nutrient environment. The overlap in δ(15)N values with multiple nitrogenous sources precluded identification. However, spatial and temporal patterns did not support stormwater as the dominant, nitrogenous source. Patterns were congruent with algal incorporation of terrestrial derived N, subjected to a high degree of biogeochemical cycling.

Algal δ15N values detect a wastewater effluent plume in nearshore and offshore surface waters and three-dimensionally model the plume across a coral reef on Maui, Hawai'i, USA.

The coral reef at Kahekili, Maui is located ~300 m south of the Lahaina Wastewater Reclamation Facility which uses four Class V injection wells to dispose of 3-5 million gallons of wastewater effluent daily. Prior research documented that the wastewater effluent percolates into the nearshore region of Kahekili. To determine if the wastewater effluent was detectable in the surface waters offshore, we used algal bioassays from the nearshore region to 100 m offshore and throughout the water column from the surface to the benthos. These algal bioassays documented that significantly more wastewater effluent was detected in the surface rather than the benthic waters and allowed us to generate a three-dimensional model of the wastewater plume in the Kahekili coastal region. Samples located over freshwater seeps had the highest δ(15)N values (~30-35‰) and the effluent was detected in surface samples 500 m south and 100 m offshore of the freshwater seeps (~8-11‰).

Using delta 15N values in algal tissue to map locations and potential sources of anthropogenic nutrient inputs on the island of Maui, Hawai'i, USA.

Macroalgal blooms of Hypnea musciformis and Ulvafasciata in coastal waters of Maui only occur in areas of substantial anthropogenic nutrient input, sources of which include wastewater effluent via injection wells, leaking cesspools and agricultural fertilizers. Algal delta(15)N signatures were used to map anthropogenic nitrogen through coastal surveys (island-wide and fine-scale) and algal deployments along nearshore and offshore gradients. Algal delta(15)N values of 9.8 per thousand and 2.0-3.5 per thousand in Waiehu and across the north-central coast, respectively, suggest that cesspool and agricultural nitrogen reached the respective adjacent coastlines. Effluent was detected in areas proximal to the Wastewater Reclamation Facilities (WWRF) operating Class V injection wells in Lahaina, Kihei and Kahului through elevated algal delta(15)N values (17.8-50.1 per thousand). From 1997 to 2008, the three WWRFs injected an estimated total volume of 193 million cubic meters (51 billion gallons) of effluent with a nitrogen mass of 1.74 million kilograms (3.84 million pounds).

The effects of top-down versus bottom-up control on benthic coral reef community structure.

While climate change and associated increases in sea surface temperature and ocean acidification, are among the most important global stressors to coral reefs, overfishing and nutrient pollution are among the most significant local threats. Here we examined the independent and interactive effects of reduced grazing pressure and nutrient enrichment using settlement tiles on a coral-dominated reef via long-term manipulative experimentation. We found that unique assemblages developed in each treatment combination confirming that both nutrients and herbivores are important drivers of reef community structure. When herbivores were removed, fleshy algae dominated, while crustose coralline algae (CCA) and coral were more abundant when herbivores were present. The effects of fertilization varied depending on herbivore treatment; without herbivores fleshy algae increased in abundance and with herbivores, CCA increased. Coral recruits only persisted in treatments exposed to grazers. Herbivore removal resulted in rapid changes in community structure while there was a lag in response to fertilization. Lastly, re-exposure of communities to natural herbivore populations caused reversals in benthic community trajectories but the effects of fertilization remained for at least 2 months. These results suggest that increasing herbivore populations on degraded reefs may be an effective strategy for restoring ecosystem structure and function and in reversing coral-algal phase-shifts but that this strategy may be most effective in the absence of other confounding disturbances such as nutrient pollution.