Herbivorous sea urchins (Echinometra mathaei) support resilience on overfished and sedimented tropical reefs.

Human impacts are dramatically changing ecological communities, motivating research on resilience. Tropical reefs are increasingly undergoing transitions to short algal turf, a successional community that mediates either recovery to coral by allowing recruitment or transitions to longer turf/macroalgae. Intense herbivory limits turf height; subsequently, overfishing erodes resilience of the desirable coral-dominated reef state. Increased sedimentation also erodes resilience through smothering and herbivory suppression. In spite of this critical role, most herbivory studies on tropical reefs focus on fishes, and the contribution of urchins remains under-studied. To test how different herbivory and sedimentation scenarios impact turf resilience, we experimentally simulated, in situ, four future overfishing scenarios derived from patterns of fish and urchin loss in other reef systems and two future sedimentation regimes. We found urchins were critical to short turf resilience, maintaining this state even with reduced fish herbivory and increased sediment. Further, urchins cleared sediment, facilitating fish herbivory. This study articulates the likelihood of increased reliance on urchins on impacted reefs in the Anthropocene.

A persistent green macroalgal mat shifts ecological functioning and composition of associated species on an Eastern Tropical Pacific coral reef.

Global evidence of phase shifts to alternate community types is of particular concern because these new communities can provide fundamentally different and often novel ecosystem functions and services compared to the original community. Shifts of a diverse range of marine communities to dominance by green macroalgal mats have occurred worldwide, making it critical to understand their emerging functions and roles. We observed a green algal mat on two reefs in the Eastern Tropical Pacific, with one persisting for >10 years on a reef with stable herbivore populations and no known sources of anthropogenic nutrients. These mats supported a more speciose macroalgal community with fewer taxa present in the adjacent coral community and facilitated growth of an associated understory macroalgal species by reducing herbivory pressure and possibly enhancing nutrient supplies within the mat community state. These results demonstrate a weakening in the processes controlling reef community structure as a result of the shift in composition associated with the macroalgal mat, creating a positive feedback supporting mat persistence. These novel ecosystem functions generated by this alternate community state illustrate the importance of further research on community shifts, which will become increasingly common in the Anthropocene.

Increased dominance of heat-tolerant symbionts creates resilient coral reefs in near-term ocean warming.

Climate change is radically altering coral reef ecosystems, mainly through increasingly frequent and severe bleaching events. Yet, some reefs have exhibited higher thermal tolerance after bleaching severely the first time. To understand changes in thermal tolerance in the eastern tropical Pacific (ETP), we compiled four decades of temperature, coral cover, coral bleaching, and mortality data, including three mass bleaching events during the 1982 to 1983, 1997 to 1998 and 2015 to 2016 El Niño heatwaves. Higher heat resistance in later bleaching events was detected in the dominant framework-building genus, Pocillopora, while other coral taxa exhibited similar susceptibility across events. Genetic analyses of Pocillopora spp. colonies and their algal symbionts (2014 to 2016) revealed that one of two Pocillopora lineages present in the region (Pocillopora "type 1") increased its association with thermotolerant algal symbionts (Durusdinium glynnii) during the 2015 to 2016 heat stress event. This lineage experienced lower bleaching and mortality compared with Pocillopora "type 3", which did not acquire D. glynnii. Under projected thermal stress, ETP reefs may be able to preserve high coral cover through the 2060s or later, mainly composed of Pocillopora colonies that associate with D. glynnii. However, although the low-diversity, high-cover reefs of the ETP could illustrate a potential functional state for some future reefs, this state may only be temporary unless global greenhouse gas emissions and resultant global warming are curtailed.

Ontogenetic Variation in Blade Toughness May Contribute to the Spread of Turbinaria ornata Across the South Pacific.

Coral reefs are shifting from coral to algal-dominated ecosystems worldwide. Recently, Turbinaria ornata, a marine alga native to coral reefs of the South Pacific, has spread in both range and habitat usage. Given dense stands of T. ornata can function as an alternative stable state on coral reefs, it is imperative to understand the factors that underlie its success. We tested the hypothesis that T. ornata demonstrates ontogenetic variation in allocation to anti-herbivore defense, specifically that blade toughness varied nonlinearly with thallus size. We quantified the relationship between T. ornata blade toughness and thallus size for individual thalli within algal stands (N = 345) on seven fringing reefs along the north shore of Moorea, French Polynesia. We found that blade toughness was greatest at intermediate sizes that typically form canopies, with overall reduced toughness in both smaller individuals that refuge within the understory and older reproductive individuals that ultimately detach and form floating rafts. We posit this variation in blade toughness reduces herbivory on the thalli that are most exposed to herbivores and may facilitate reproduction in dispersing stages, both of which may aid the proliferation of T. ornata.

Flip it and reverse it: Reasonable changes in designated controls can flip synergisms to antagonisms.

Ecological systems are subjected to multiple stressors that can interact in complex ways resulting in "ecological surprises". We examine the pivotal role of 'control' assignment in the categorization of stressors into five classes: additive, +synergistic, -synergistic, +antagonistic, and -antagonistic. We demonstrate if an alternate treatment can reasonably be considered the experimental control, nonlinear interaction classifications change, both in sign (+/-) and in direction (synergistic/antagonistic). Further, switching of interaction classifications is not predictable as changing control can result in multiple possible alternate nonlinear classifications. To explore the magnitude of this problem, we evaluate publications gathered for a recent meta-analysis to 1) explore rationales for choice of controls and 2) quantify how frequently it is reasonable to reassign the control. We found controls were designated with a variety of implicit and explicit justifications, with two overall rationales: 1) controls based on 'natural' conditions (historic, current, or future); 2) controls based on direction of impact, such that stressors always have negative impacts. We reasoned that control re-assignment was justified if an alternate treatment met one of these rationales. Of the 844 interactions classified in the meta-analysis, we determined >95% could be reassigned. Based on these findings, we recommend a new approach to meta-analyses, where the 'control' is strictly and consistently defined by the authors of the meta-analysis. These controls should be based on their broader question, rather than following the common practice of defaulting to controls assigned by the authors of each study, as we found these rationales vary broadly based on the specific questions of each study. Consistent control designation within the ecological or toxicological framework of each meta-analysis may provide deeper and more consistent insight into the nature of interactive effects between multiple stressors. Gaining this insight is crucial because stressor interactions are certain to increase in the Anthropocene.

Storms may disrupt top-down control of algal turf on fringing reefs.

Storms strongly affect coral reefs; one unstudied but potentially important outcome may be a decrease in herbivory, presumably through changes to freshwater, sediment and nutrient influx. Algal turfs are sensitive early indicators of reef condition, and experimental evidence demonstrates low sediment loads and strong herbivory maintain short, healthy turf. While unknown, storms likely disrupt these controlling forces. We have observed storms that generate frequent, visible sediment plumes in Moorea, French Polynesia. To evaluate the effects of storms on herbivory, we conducted a set of field experiments manipulating herbivore access to naturally occurring turf under three rainfall regimes: no rain, light rain, and heavy rain that generated a plume event. We found strong effects of herbivores except following the storm, indicating disruption of typically strong top-down control by herbivores on algal turfs. Further research into the underlying mechanisms is critical as storm intensities and watershed development increase in many tropical regions.

Selective consumption of macroalgal species by herbivorous fishes suggests reduced functional complementarity on a fringing reef in Moorea, French Polynesia.

Worldwide, many coral reefs are at risk of shifting to degraded algal-dominated states, due to compromised ecological conditions. Functional diversity of herbivorous fishes maintains coral reef health and promotes reef resilience to disturbances. Given previous evidence, it appears the functional roles of herbivorous fishes differ across geographical locations, indicating a need for further assessment of macroalgal consumption by herbivorous fishes. We assessed functional diversity by examining foraging behavior of herbivorous fish species on macroalgae on a fringing reef in Moorea, French Polynesia. We video-recorded choice experiments containing seven common macroalgae and used Strauss' linear resource selection index to determine macroalgal selectivities. We used cluster analysis to identify any distinct groups within herbivorous fish species, given the macroalgal species they targeted, and fitted generalized linear mixed-effects models to identify factors that best predicted the number of bites taken on macroalgae. Seven species from 3 fish families/tribes took a total of 956 bites. Fish species differed in their selectivity with some species (Naso lituratus, N. unicornis, Calatomus carolinus) strongly preferring one or two macroalgal species, while other fish species (Acanthurus nigrofuscus, Ctenochaetus striatus, Chlorurus sordidus, Balistapus undulatus) were less selective. This resulted in fish species forming two clusters. Only 3 of 7 macroalgae were preferred by any fish species, with two fish species both preferring the same two macroalgae. The limited differences in fish species' preferences for different macroalgae suggests limited functional complementarity. Two models (macroalgal species identity+fish functional group, macroalgal species identity+fish species) best predicted the number of bites taken on macroalgae compared to models incorporating only a single explanatory factor or fish family. In the context of this Moorean fringing reef, there is greater functional redundancy than complementarity of herbivorous fishes consuming macroalgae, and the fishes grouped together according to their relative selectivity. We observed fish species that are not classified as browsers consuming macroalgae, suggesting diets of herbivorous fishes may be broader than previously thought. Finally, we observed macroalgal selectivities and consumption that differed from previous studies for the same fish species. Our results contribute to the understanding of functional diversity of herbivorous fishes across coral reefs, and also highlight the need for additional research to further elucidate the role of context and functional diversity of herbivorous fishes consuming macroalgae on coral reefs.

TESTING THE CONCEPTUAL AND OPERATIONAL UNDERPINNINGS OF HERBIVORY ASSAYS: DOES VARIATION IN PREDICTABILITY OF RESOURCES, ASSAY DESIGN, AND DEPLOYMENT METHOD AFFECT OUTCOMES?

Herbivory assays are a valuable tool used by ecologists to understand many of the patterns and processes affecting herbivory, a widely recognized driving force in marine communities. However, methods vary substantially among studies in both design and operation, and the effect of these differences has yet to be evaluated. We assessed the effects of several key components of assay design on estimates of herbivory to offer four recommendations. First, we found assays out-planted on sequential days in both predictable and random locations within a 60m2 site experienced temporal increases in herbivory by an increasingly diverse assemblage of fishes. Thus, we strongly advise against placing herbivory assays in the same site over a series of days. Second, we found while the amount of biomass consumed in assays was density dependent, the percent loss was not. Thus, we recommend researchers report percent consumption because this metric is robust to differences in biomass offered and will facilitate comparisons across studies. Third, we found associational effects, where proximity of species of differing palatabilities impacted estimates of herbivory rate on one or both species, but these impacts were not consistent across species or sites. Thus, we recommend the effect of association be directly tested for multi specie herbivory assays. Fourth, we found no effect of attachment method on estimates of herbivory rate and recommend researchers continue to use the attachment method in which they are most confident. We hope our experimental results prove useful in the future when designing, conducting, and interpreting herbivory assays.

Epibionts on Turbinaria ornata, a secondary foundational macroalga on coral reefs, provide diverse trophic support to fishes.

Worldwide, many coral reef ecosystems have shifted from coral to algal dominance, yet the ecological function of these emergent communities remains relatively unknown. Turbinaria ornata, a macroalga with a rapidly expanding range in the South Pacific, forms dense stands on hard substrate, likely providing ecological services unique from corals. While generally unpalatable, T. ornata can function as a secondary foundation species and hosts an epibiont community that may provide overlooked trophic resources in phase shifted reef ecosystems. Results from video recorded field experiments designed to quantify consumer pressure on T. ornata epibionts showed that both consumer pressure and epibiont cover increased with thallus size. Additionally, most fish species, including herbivores, omnivores, and detritivores, exhibited higher bite rates on thalli with epibionts compared to thali with epibionts experimentally removed. Juvenile parrotfishes were responsible for 50% of total bites recorded and also had the highest bite rates. Results indicate that epibionts, particularly on large T. ornata, are a food resource for a diversity of fishes, representing a previously undescribed function of this macroalga in coral reef ecosystems. Exploring the functions of macroalgal dominated reef communities will be increasingly important as reefs continue to phase shift toward macroalgal dominance in the Anthropocene.

State of corals and coral reefs of the Galápagos Islands (Ecuador): Past, present and future.

Coral populations and structural coral reefs have undergone severe reductions and losses respectively over large parts of the Galápagos Islands during and following the 1982-83 El Niño event. Coral tissue loss amounted to 95% across the Archipelago. Also at that time, all coral reefs in the central and southern islands disappeared following severe degradation and eventual collapse due primarily to intense bioerosion and low recruitment. Six sites in the southern islands have demonstrated low to moderate coral community (scattered colonies, but no carbonate framework) recovery. The iconic pocilloporid reef at Devil's Crown (Floreana Island) experienced recovery to 2007, then severe mortality during a La Niña cooling event, and is again (as of 2017) undergoing rapid recovery. Notable recovery has occurred at the central (Marchena) and northern islands (Darwin and Wolf). Of the 17 structural reefs first observed in the mid-1970s, the single surviving reef (Wellington Reef) at Darwin Island remains in a positive growth mode. The remainder either degraded to a coral community or was lost. Retrospective analyses of the age structure of corals killed in 1983, and isotopic signatures of the skeletal growth record of massive corals suggest the occurrence of robust coral populations during at least a 500-year period before 1983. The greatest potential threats to the recovery and persistence of coral reefs include: ocean warming and acidification, bioerosion, coral diseases, human population growth (increasing numbers of residents and tourists), overfishing, invasive species, pollution, and habitat destruction. Such a diverse spectrum of disturbances, acting alone or in combination, are expected to continue to cause local and archipelago-wide mortality and degradation of the coral reef ecosystem.

Rapid recovery of a coral dominated Eastern Tropical Pacific reef after experimentally produced anthropogenic disturbance.

Local anthropogenic stressors such as overfishing, nutrient enrichment and increased sediment loading have been shown to push coral reefs toward greater dominance by algae. In a few cases this shift has been temporary, with the ability to recover to a healthy coral-dominated community after disturbance, suggesting some systems have considerable resilience. However, an understanding of the circumstances under which reefs may recover is only beginning to emerge. We monitored recovery of a coral-dominated reef in the Eastern Tropical Pacific (ETP) after cessation of a ∼6 month multiple stressor experiment (with herbivore exclosure, nutrient addition, and sediment addition). We observed substantial recovery from small-scale disturbances, though there were differences in both the extent and temporal dynamics of recovery between treatments. Plots that had been caged showed the largest recovery in absolute terms and recovery was quite rapid, while nutrient and sediment addition plots were slower to recover. We also observed different recovery patterns depending on the type of algae that replaced coral during or after disturbances. Macroalgae that established during manipulation were almost completely removed within 2 weeks, revealing that a significant proportion had covered still-living coral. Turf algae persisted longer, but were almost completely replaced by regenerating coral within 18 months. Very little crustose coralline algae were apparent during manipulations, but coverage did increase during recovery. This rapid recovery of corals after simulated anthropogenic disturbance to ETP reefs underscores the value of management of local stressors for short-term recovery and perhaps as a buffer for longer-term global stressors.

Empirical data demonstrates risk-tradeoffs between landscapes for herbivorous fish may promote reef resilience.

Herbivores balance resource requirements with predation risk, which can differ among landscapes; hence, landscape can shape these trade-offs, influencing herbivore distribution and behavior. While this paradigm has been well established on coral-dominated reefs, tropical reefs worldwide are shifting to algal dominance. If herbivores avoid algae due to higher risk and forage in coral, these algal states may be stabilized. However, if herbivores forage more in resource-rich algal states, this may promote coral recovery. We assessed the distribution and behavior of herbivorous fishes in Moorea, French Polynesia in coral and algal turf-dominated fringing reef sites. Acanthuridae were more abundant in coral states and Labridae, tribe Scarinae, in algal turf states, though total fish abundances were equivalent in the two states. Fish in both families spent more time feeding in algal states and hiding/swimming in coral states. Thus, behavior reflects the trade-off between resource acquisition and refuge in these two landscapes and may promote recovery to coral.

Why species matter: an experimental assessment of assumptions and predictive ability of two functional-group models.

Community ecologists use functional groups based on the rarely tested assumption that within-group responses to ecological processes are similar and thus members are functionally equivalent. However, recent research suggests that functional equivalency may break down with human impacts. We tested the equivalency assumption and model predictions of responses to simulated human alterations in nutrients and large herbivores for two models of coral reef algae, the Relative Dominance Model (RDM) and the Functional Group Model (FGM). Results of both mesocosm and field experiments using assembled communities were compared to model predictions, and within- and between-group variability were assessed. Both models' predictions of group response to herbivory matched experimental outcomes, but only the RDM predicted response to nutrients. However, within-group variability was dramatic, because the RDM grouped species with opposite responses to herbivory and the FGM grouped species with unique responses to nutrients. These heterogeneous responses resulted in loss of information and masked strong interactions between herbivory and nutrients that were not included in the models. As humans continue to impact major ecological processes in ecosystems globally, we postulate that functional-group models may need to be reformulated to account for shifting baselines.

How much is too much? Identifying benchmarks of adverse effects of macroalgae on the macrofauna in intertidal flats.

Eutrophication, defined as the accumulation of organic matter typically in response to anthropogenically enhanced nutrient inputs, often takes the form of macroalgal blooms in shallow estuaries and causes a cascade of adverse ecosystem effects. Confidence in the use of macroalgae as an indicator of eutrophication in estuaries is limited by the lack of quantitative data on thresholds of adverse effects. Field experiments can provide "benchmarks" of no effect or adverse effects that can be used to validate thresholds derived statistically from field data. To determine a benchmark of adverse effects of macroalgal abundance on macrobenthic faunal communities in intertidal flats, experiments were conducted in two sites in Bodega Harbor (BOD) and two sites in Upper Newport Bay (UNB), California, USA. At each site, 24 cages maintained six treatments of macroalgae for eight weeks, with mat depths of 0, 1.0, 1.5, 2.5, 3.5, and 5.0 cm composed mostly of bloom-forming green macroalgae in the genus Ulva. Every two weeks, cores of sediment (10 cm deep) were collected, and macrofauna were quantified. Mats 1 cm deep, equivalent to a biomass of 110-120 g dry mass (dm)/m2 or 840-930 g wet mass/m2, resulted in the reduction of macrofaunal abundance by at least 67% and species richness by at least 19% within two weeks at three of four sites. Loss was attributed to the decline of key functional groups. Surface-deposit feeders were eliminated from one site at BOD within four weeks and at one site in UNB within six weeks, while 1-cm mats negatively affected suspension feeders and herbivores in the second site at BOD. In contrast, the other site at UNB was not affected by macroalgal treatment, likely due to an initial community composed of a high proportion of subsurface-deposit feeders tolerant of stressful environments. Macroalgal abundances as low as 110-120 g dm/m2 had significant and rapid negative effects on macrobenthic invertebrates, providing a clear benchmark of adverse effects of macroalgal blooms on macrofaunal abundance and community structure, two indicators of ecosystem health. This information can inform the establishment of appropriate metrics for macroalgal abundance in eutrophic estuaries.

RAPID NITRATE UPTAKE RATES AND LARGE SHORT-TERM STORAGE CAPACITIES MAY EXPLAIN WHY OPPORTUNISTIC GREEN MACROALGAE DOMINATE SHALLOW EUTROPHIC ESTUARIES1.

We quantified the effects of initial macroalgal tissue nitrogen (N) status (depleted and enriched) and varying pulses of nitrate (NO3- ) concentration on uptake and storage of nitrogen in Ulva intestinalis L. and Ulva expansa (Setch.) Setch. et N. L. Gardner using mesocosms modeling shallow coastal estuaries in Mediterranean climates. Uptake of NO3- (µmol · g dry weight [dwt]-1 · h-1 ) was measured as loss from the water after 1, 2, 4, 8, 12, and 24 h and storage as total tissue nitrogen (% dwt) and nitrate (ppm). Both species of algae exhibited a high affinity for NO3- across all N pulses and initial tissue contents. There was greater NO3- removal from the water for depleted than enriched algae across all time intervals. In the low-N-pulse treatment, U. intestinalis and U. expansa removed all measurable NO3- within 8 and 12 h, respectively, and in the medium and high treatments, removal was high and then decreased over time. Maximum mean uptake rates of nitrate were greater for U. expansa (∼300 µmol · g dwt-1 · h-1 ) than U. intestinalis (∼100 µmol · g dwt-1 · h-1 ); however, uptake rates were highly variable over time. Overall, U. expansa uptake rates were double those of U. intestinalis. Maximum tissue NO3- for U. expansa was >1,000 ppm, five times that of U. intestinalis, suggesting that U. expansa has a greater storage capacity in this cellular pool. These results showed that opportunistic green algae with differing tissue nutrient histories were able to efficiently remove nitrate from the water across a wide range of N pulses; thus, both are highly adapted to proliferate in estuarine environments with pulsed nutrient supplies.

Does use of tropical beaches by tourists and island residents result in damage to fringing coral reefs? A case study in Moorea French Polynesia.

Although coral reefs worldwide are subject to increasing global threats, humans also impact coral reefs directly through localized activities such as snorkeling, kayaking and fishing. We investigated five sites on the northern shore of Moorea, French Polynesia, and quantified the number of visitors on the beach and in shallow water. In field surveys, we measured total coral cover and colony sizes of two common genera, Porites and Acropora, a massive and branching morphology, respectively. One site, which hosted over an order of magnitude more people than the other four, had significantly less total coral cover and supported very little branching Acropora. In addition, size frequency distributions of both the branching and massive genera were skewed toward smaller colony sizes at the high use site. Our results demonstrated that the use of tropical beaches may result in less coral cover, with branching colonies rare and small.

Spatial refuges and associational defenses promote harmful blooms of the alga Caulerpa sertularioides onto coral reefs.

Extreme population fluctuations, or outbreaks, are driven by interacting processes that are often more complex than isolated changes in consumer or resource control. Blooms of the macroalga Caulerpa sertularioides in the eastern tropical Pacific overgrew and killed reef-building corals, with blooms onto reefs corresponding to cool La Niña phases of inter-decadal fluctuations of the El Niño-Southern Oscillation. We quantified factors responsible for the maintenance of C. sertularioides patches in off-reef areas, namely an associational mutualism with an epiphytic cyanobacteria (Lyngbya majuscula), coupled with spatial refuges at the scales of individual thalli and habitat. Maintenance of near reef algal populations with a strong response to nutrient addition showed that these populations were primed to bloom onto reefs in response to enhanced nutrient delivery, such as those potentially associated with La Niña conditions. However, our experiments demonstrated that no single factor related to consumer or resource control was likely to stimulate bloom formation in isolation. Rather, we propose a novel model of reef bloom formation where off-reef blooms are sustained by processes reducing consumer control, and then bloom onto reefs through an interaction between increased allochthonous nutrient input and an uncoupling of consumer control by an association with epiphytic cyanobacteria.

Decoupling of nutrient and grazer impacts on a benthic estuarine diatom assemblage.

Strong interactions between top-down (consumptive) and bottom-up (resource supply) trophic factors occur in many aquatic communities, but these forces can act independently in some microphytobenthic communities. Within benthic estuarine diatom assemblages, the dynamics of these interactions and how they vary with abiotic environmental conditions are not well understood. We conducted a field experiment at two sites with varying habitat characteristics to investigate the interactive effects of grazers and nutrients on benthic estuarine diatoms. We crossed snail (Cerithidea californica) and nutrient (nitrogen and phosphorus) addition treatments in enclosures on a restored tidal sandflat and a reference tidal mudflat in Mugu Lagoon, southern California. We repeated the study in summer 2000 and spring 2001 to assess temporal variation in the interactions. Snails caused a large decrease in diatom relative abundance and biomass (estimated as surface area); nutrients increased diatom abundance but did not alter diatom biomass. Snails and nutrients both reduced average diatom length, although the nutrient effect was weaker and temporally variable, occurring in the reference mudflat in the spring. There were few interactions between snail and nutrient addition treatments, suggesting that links between top-down and bottom-up forces on the diatom community were weak. There were no consistent differences in diatom assemblage characteristics between the two study sites, despite marked differences in sediment grain size and other abiotic characteristics between the sites. The strong diatom response to herbivores and weaker responses to enrichment differed from the previous studies where cyanobacteria increased in response to nutrient enrichment, further dissolving the "black box" perception of microphytobenthic communities.

The impacts of human visitation on mussel bed communities along the California coast: are regulatory marine reserves effective in protecting these communities?

Rocky intertidal habitats frequently are used by humans for recreational, educational, and subsistence-harvesting purposes, with intertidal populations damaged by visitation activities such as extraction, trampling, and handling. California Marine Managed Areas, particularly regulatory marine reserves (MRs), were established to provide legal protection and enhancement of coastal resources and include prohibitions on harvesting intertidal populations. However, the effectiveness of MRs is unclear as enforcement of no-take laws is weak and no regulations protect intertidal species from other detrimental visitor impacts such as trampling. The goal of this study was two-fold: (1) to determine impacts from human visitation on California mussel populations (Mytilus californianus) and mussel bed community diversity; and (2) to investigate the effectiveness of regulatory MRs in reducing visitor impacts on these populations. Surveys of mussel populations and bed-associated diversity were compared: (1) at sites subjected to either high or low levels of human use, and (2) at sites either unprotected or with regulatory protection banning collecting. At sites subjected to higher levels of human visitation, mussel populations were significantly lower than low-use sites. Comparisons of mussel populations inside and outside of regulatory MRs revealed no consistent pattern suggesting that California no-take regulatory reserves may have limited effectiveness in protecting mussel communities. In areas where many people visit intertidal habitats for purposes other than collecting, many organisms will be affected by trampling, turning of rocks, and handling. In these cases, effective protection of rocky intertidal communities requires an approach that goes beyond the singular focus on collecting to reduce the full suite of impacts.

Macroalgal bioindicators (growth, tissue N, delta(15)N) detect nutrient enrichment from shrimp farm effluent entering Opunohu Bay, Moorea, French Polynesia.

Nutrient enrichment from shrimp aquaculture poses an increasing environmental threat due to the industry's projected rapid growth and unsustainable management practices. Traditional methods to monitor impacts emphasize water quality sampling; however, there are many advantages to bioindicators, especially in developing countries. We investigated the usefulness of three bioindicators -- growth, tissue nitrogen content and nitrogen stable isotope signature (delta(15)N) -- in the tropical red macroalga Acanthophora spicifera. Algae were collected, cultured, and deployed in a spatial array around the outflow from a shrimp farm in Moorea, French Polynesia, to detect nitrogenous wastes. All three parameters were highest adjacent to the shrimp farm indicating nutrient enrichment, and delta(15)N values confirmed the shrimp farm as the dominant nutrient source (5.63-5.96 per thousand). Isotope ratios proved the most sensitive indicator, as delta(15)N signatures were detected at the most distant sites tested, confirming their usefulness in tracing nutrients and mapping the spatial extent of enrichment.