Benthic cover data from six sites 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

Benthic cover data from six sites 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 benthic cover of the 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.

Benthic cover: The benthic cover of the six study sites was surveyed along 30-m long transects running parallel to the shore in each MPA and non-MPA. Along each transect, photographs were taken from 0.5 m above the bottom every 2 m along each transect (i.e., 16 photos per transect), so that consecutive photographs did not overlap. The area of each photograph was about 25 cm x 30 cm, therefore sampled area per transect was about 1.2m2. Photos were analyzed for percentage cover of corals and macrophytes using CPC with Excel extensions; the program randomly placed 20 points on each photo, and we identified the organism beneath each point. Organisms were further classified into four main categories: scleractinian corals, macroalgae, epilithic algal matrix [the EAM sensu 64], and others.

A total of 273 transects were conducted: 87 in Votua (48 in the MPA and 39 in the non-MPA), 94 in Vatu-o-lalai (44 in the MPA and 50 in the non-MPA), and 87 in Namada (41 in the MPA and 51 in the non-MPA). Transects were conducted from December 2010 to January 2011 and from December 2011 to January 2012, and there was no spatial overlap between transects even between different sampled year periods. On each sampling day, four to six transects were deployed on the reef parallel to the shoreline. To ensure that transects were independent and non-overlapping, they were separated by a minimum of 15 m, and the ends of each transect were marked with small surface buoys. Additionally, after conducting every 4–6 transects, the snorkeler swam approx. 25 m towards the reef crest to start a new set of transects, so sampling was widespread within each area.

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), 12.96 KB)
Primary data file for dataset ID 674104

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siteSite name unitless
statusStatus of area where sampling was done; Marine Protected Area (MPA) or non-MPA (NON) unitless
transectTransect number unitless
year(A) Sampling took place between December 2010 and January 2011; (B) Sampling took place between December 2011 and January 2012 unitless
coralPercent cover of coral centimeters
maPercent cover of macroalgae centimeters
eamPercent cover of epilithic algal matrix centimeters
otherPercent cover of other categories centimeters

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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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