Biomass and density data from in and outside of an MPA in Viti Levu, Fiji from 2010-2012 (Killer Seaweeds project)

Data Type: Other Field Results
Version: 1
Version Date: 2017-01-06

» Killer Seaweeds: Allelopathy against Fijian Corals (Killer Seaweeds)
Hay, MarkGeorgia Institute of Technology (GA Tech)Principal Investigator
Ake, HannahWoods Hole Oceanographic Institution (WHOI BCO-DMO)BCO-DMO Data Manager

Biomass and density data from in and outside of an MPA in Viti Levu, Fiji from 2010-2012 (Killer Seaweeds project)


Temporal Extent: 2010 - 2012

Dataset Description

Fiji MPA vs non-MPA corals, fishes, and herbivory. This dataset contains information on the biomass and density at six study sites.

Methods & Sampling

The study was conducted from November 2010 through February 2011 and between November 2011 and January 2012 on shallow (~1 m below the surface at low tide, equal or shallower than 2 m at high tide), intertidal fringing reefs platforms (up to 800-m wide) along the Coral Coast (18º 13.05’S, 177º 42.97’E) of Viti Levu, Fiji’s main island. Many of the owners of traditional fishing rights along the Coral Coast have established small, customary no-take MPAs to improve and sustain their adjacent fishing grounds. The MPAs in this region are delimited by surface markings and enforced by local villagers, and they have been closed to all fishing activities since their inception (about 10 years). The only exception to this closure was a small experimental hook and line fishing research project that was conducted in the MPAs of Votua and Namada. In the non-MPAs, the main fishing targets are species of Acanthuridae (Nasinae), Epinephelidae, Labridae, Mullidae, and Lutjanidae. Permission for the research was granted by the Fijian Ministry of Education, National Heritage, Culture & Arts, Youth & Sports, which is authorized to approve field studies in Fijian waters. No animal collection or experimental procedures involving animals were conducted during the study, and no endangered species were recorded during our assessments. 

To assess the effects of MPAs on fish assemblages, fish feeding group composition, herbivory rates, benthic cover, and coral recruit density, we compared three spatially paired MPA and adjacent, fished, areas (non-MPAs) associated with the villages of Votua, Vatu-o-lalai and Namada.  Comparisons of fish assemblages inside and outside of closures are widely used for determining the effects of reserves, but it should be acknowledged that this approach does not reveal the state of an MPA relative to an undisturbed baseline. 

The studied MPAs were established in 2002 (Vatu-o-lalai, Namada) and 2003 (Votua), and shortly after establishment, coral cover was low (~7%), and macroalgal cover was high (~35–45%) in both the MPAs and non-MPAs. All surveys and assays were conducted during the same season (austral summer) to minimize seasonal variation in sampling. The reef extends approx. 1 km from shore within each MPA and non-MPA, and all data were collected between 30 and 700 m of the shore (i.e., shoreward of the reef crest) parallel to the shoreline.

Fish assemblages: Underwater visual censuses (UVC) were used to assess fish assemblages in MPAs and non-MPAs at the three village sites. Underwater visibility at all study sites (> 15 m) was appropriate for the use of UVC, but due to the visual limitations of this method, we did not consider cryptic species or species with a maximum total length < 5 cm. During our surveys, we categorized species into two major categories (Herbivores and Non-herbivores) that were subdivided into ten sub-categories. Herbivores include the main roving nominally herbivorous fish clades, which play an important role in the control of benthic algae, and these species were further divided into four sub-categories (browsers, grazers, scrapers, and excavators) according to diet, feeding mode, and impact on the benthos. The category of Non-herbivores includes all species that feed on other, non-algal resources.

Separate 30m x 4m belt transects were performed for Herbivores and Non-herbivores. While simultaneously deploying the transect line, a snorkeler recorded all non-cryptic fishes (either Herbivores or Non-herbivores) within 2 m of either side of the transect. Individual fish were identified to species and placed into 5-cm (total length) size classes, and the lengths were converted to biomass using established length-weight relationships. 

Transects were conducted in each area within 2 h of high tide (approx. 1.5 m depth) and were equally distributed between the two sampling periods (Dec 2010–Jan 2011 and Dec 2011–Jan 2012), the months within each sampled year (December and January of each year). On each sampling day, four to six transects were deployed on the reef parallel to the shoreline, with a minimum of 10m between adjacent transects. To ensure that transects were independent and non-overlapping, small numbered surface floats were placed at the start and end of each transect, and were left in position during all sampling. Care was taken to avoid re-counting fishes that left and subsequently re-entered the transect areas. The initial starting point of the transects for each day was selected based on a map of the study sites with two constraints: (1) as to a minimum distance from shore (at least 30 m), and a minimum distance from the MPA boundaries (150 m). 

Density of coral recruits: To assess the effect of the MPAs on the replenishment of coral populations, the density of coral recruits on natural reef substrata was assessed at night during January and February 2011 using a fluorescence technique. Coral recruit counts were conducted along a series of 50-m long transects positioned parallel to shore, and a total of 18 transects (n = 9 MPA; n = 9 non-MPA) were performed at each of the three village sites (n = 54 transects total). Ten 25 cm x 30 cm rectangular quadrats were randomly placed along each transect with a minimum distance of 2 m between quadrats (540 plots total). Once a quadrat was deployed on the substratum, all coral recruits within the borders of the quadrat were counted. 

Data Processing Description

We used Generalized linear mixed models (GLMM) implemented under a Bayesian framework to test the effect of protection status (MPA vs. non-MPA) on the abundance and biomass of each subcategory of Herbivores and Non-herbivores at the three village sites. We used the same approach to test the effect of protection status on the number of individuals, biomass and diversity of species in the observed feeding groups. The models have a hierarchical structure where the protection status is nested within site. For abundance and biomass of Herbivores and Non-herbivores, we used a muti-response models where each subcategory is a separate response variable. Because we have several samples for the same site, transect and sampling day were included as random factors. For the richness and abundance model, we used a Poisson error structure given the nature of the data. For biomass and diversity data we used a Gaussian error structure. We performed separate analyses for Herbivores and Non-herbivores. We compared model fit against a benchmark model in which protection status was not included as a fixed effect using the Deviance information criterion (DIC). The MCMC used to sample the posterior distributions of effect sizes ran for 106 iterations and was sampled every 100 iterations (thinning = 100) after burn-in (5×105). We considered effect size significant when the 95% credible interval of the estimated posterior distributions of parameters did not include 0. We monitored chain mixing by checking the effective sample sizes (ESS) for fixed and random effects. We used inverse gamma priors for variance components. Exploratory analyses indicate that estimates for fixed effects were robust to prior selection. Outliers were removed prior to the GLMM analyses to reduce overdispersion, although analyses with and without the outliers yielded qualitatively similar results. We used the R package MCMCGLMM for all analyses based on GLMMs.

We compared benthic cover between MPAs and non-MPAs using three-way ANOVA, with village site (Votua, Vatu-o-lalai, and Namada), status (MPA and non-MPA) and year (2010/2011 and 2011/2012) as fixed factors. Separate ANOVAs were used to compare the percentage cover of four different substratum types (scleractinian corals, macroalgae, epilithic algal turfs and others). Benthic cover data were arcsine-transformed, and fish density and biomass data were log-transformed to meet assumptions of normality (frequency histograms). When differences were significant, the test was followed by specific planned comparisons between paired treatments (MPA vs non-MPA) at each village site. P-values were adjusted with the Holm-Sídák method, in which the adjusted p-value is equal to , where k refers to the number of comparisons.

The rates of grazing and browsing and the density of coral recruits were compared between MPAs and adjacent non-MPAs using two-way ANOVA with status (MPA and non-MPA) and village site (Votua, Vatu-o-lalai, and Namada) as fixed factors. Separate analyses were used to compare (1) parrotfish grazing rates, (2) macroalgal browsing rates, and (3) the number of coral recruits per quadrat (log-transformed). Holm-Sídák-adjusted paired comparisons were also used when differences were significant. ANOVAs for benthic cover, grazing and browsing rates, and density of coral recruits, as well as all graph plots in this manuscript, were programmed in R 3.0.1 using base package functions.

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Data Files

(Comma Separated Values (.csv), 81.95 KB)
Primary data file for dataset ID 674116

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typeCategory of fish assemblage sampled; Herbivores include the main roving nominally herbivorous fish clades; non-herbivores includes all species that feed on other, non-algal resources. unitless
siteSite name unitless
statusStatus of area where sampling was done; Marine Protected Area (MPA) or non-MPA (NON) unitless
minorSpecies were further divided into four sub-categories (browsers grazers scrapers and excavators) according to diet feeding mode and impact on the benthos. unitless
year(A) Sampling took place between December 2010 and January 2011; (B) Sampling took place between December 2011 and January 2012 unitless
dayDay that sampling took place within each years sampling event. A B and C all took place in during year A (Dec 2010 - Jan 2011); D E and F all took place during year B (Dec 2011 - Jan 2012) unitless
transectTransect number unitless
biomassBiomass of fish observed centimeters
abundanceAbundance of each species observed count

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Start Date
End Date
Studies for this deployment were conducted: November 2010 through February 2011 and between November 2011 and January 2012 on shallow (~1 m below the surface at low tide, equal or shallower than 2 m at high tide), intertidal fringing reefs platforms in Villages of Votua, Vatu-o-lalai and Namada, Coral Coast Viti Levu, Fiji. May–December 2011 on an approximately1.5-2.5 m deep reef flat within a no-take marine reserve at Votua Village, Viti Levu, Fiji.

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Project Information

Killer Seaweeds: Allelopathy against Fijian Corals (Killer Seaweeds)

Coverage: Viti Levu, Fiji (18º13.049’S, 177º42.968’E)

Extracted from the NSF award abstract:

Coral reefs are in dramatic global decline, with reefs commonly converting from species-rich and topographically-complex communities dominated by corals to species- poor and topographically-simplified communities dominated by seaweeds. These phase-shifts result in fundamental loss of ecosystem function. Despite debate about whether coral-to-algal transitions are commonly a primary cause, or simply a consequence, of coral mortality, rigorous field investigation of seaweed-coral competition has received limited attention. There is limited information on how the outcome of seaweed-coral competition varies among species or the relative importance of different competitive mechanisms in facilitating seaweed dominance. In an effort to address this topic, the PI will conduct field experiments in the tropical South Pacific (Fiji) to determine the effects of seaweeds on corals when in direct contact, which seaweeds are most damaging to corals, the role allelopathic lipids that are transferred via contact in producing these effects, the identity and surface concentrations of these metabolites, and the dynamic nature of seaweed metabolite production and coral response following contact. The herbivorous fishes most responsible for controlling allelopathic seaweeds will be identified, the roles of seaweed metabolites in allelopathy vs herbivore deterrence will be studied, and the potential for better managing and conserving critical reef herbivores so as to slow or reverse conversion of coral reef to seaweed meadows will be examined.

Preliminary results indicate that seaweeds may commonly damage corals via lipid- soluble allelochemicals. Such chemically-mediated interactions could kill or damage adult corals and produce the suppression of coral fecundity and recruitment noted by previous investigators and could precipitate positive feedback mechanisms making reef recovery increasingly unlikely as seaweed abundance increases. Chemically-mediated seaweed-coral competition may play a critical role in the degradation of present-day coral reefs. Increasing information on which seaweeds are most aggressive to corals and which herbivores best limit these seaweeds may prove useful in better managing reefs to facilitate resilience and possible recovery despite threats of global-scale stresses. Fiji is well positioned to rapidly use findings from this project for better management of reef resources because it has already erected >260 MPAs, Fijian villagers have already bought-in to the value of MPAs, and the Fiji Locally-Managed Marine Area (FLMMA) Network is well organized to get information to villagers in a culturally sensitive and useful manner.

The broader impacts of this project are far reaching. The project provides training opportunities for 2-2.5 Ph.D students and 1 undergraduate student each year in the interdisciplinary areas of marine ecology, marine conservation, and marine chemical ecology. Findings from this project will be immediately integrated into classes at Ga Tech and made available throughout Fiji via a foundation and web site that have already set-up to support marine conservation efforts in Fiji and marine education efforts both within Fiji and internationally. Business and community leaders from Atlanta (via Rotary International Service efforts) have been recruited to help organize and fund community service and outreach projects in Fiji -- several of which are likely to involve marine conservation and education based in part on these efforts there. Media outlets (National Geographic, NPR, Animal Planet, Audubon Magazine, etc.) and local Rotary clubs will be used to better disseminate these discoveries to the public.


Rasher DB, Stout EP, Engel S, Kubanek J, and ME Hay. "Macroalgal terpenes function as allelopathic agents against reef corals", Proceedings of the National Academy of Sciences, v. 108, 2011, p. 17726.

Beattie AJ, ME Hay, B Magnusson, R de Nys, J Smeathers, JFV Vincent. "Ecology and bioprospecting," Austral Ecology, v.36, 2011, p. 341.

Rasher DB and ME Hay. "Seaweed allelopathy degrades the resilience and function of coral reefs," Communicative and Integrative Biology, v.3, 2010.

Hay ME, Rasher DB. "Corals in crisis," The Scientist, v.24, 2010, p. 42.

Hay ME and DB Rasher. "Coral reefs in crisis: reversing the biotic death spiral," Faculty 1000 Biology Reports 2010, v.2, 2010.

Rasher DB and ME Hay. "Chemically rich seaweeds poison corals when not controlled by herbivores", Proceedings of the National Academy of Sciences, v.107, 2010, p. 9683.

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Funding SourceAward
NSF Division of Ocean Sciences (NSF OCE)
National Institutes of Health (NIH)

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