Scientific Papers and Reports

This is an annotated list of the published papers and reports that have included REEF data. The list is in chronological order. Papers that are available for viewing in .pdf format are noted.

Also see the Projects page for links to additional reports.

Green, S.J., N.K. Dulvy, A.L. M. Brooks, J.L. Akins, A.B. Cooper, S. Miller, and I.M. Côté. 2014. Linking removal targets to the ecological effects of invaders: a predictive model and field test.

Ecological Applications. 24: 1311–1322

The study, conducted by Dr. Stephanie Green (OSU/REEF), Lad Akins (REEF), and others, confirms for the first time that controlling lionfish populations in the western Atlantic Ocean can pave the way for a recovery of native fish. Even if it's one speared fish at a time, data are showing that removals can be effective. And not every lionfish need be removed…the research findings document that reducing lionfish numbers by specified amounts will allow a rapid recovery of native fish biomass. Over 18 months, the biomass of native prey fishes increased an average 50-70% on reefs where lionfish numbers were suppressed below target levels predicted to cause prey depletion. On reefs where lionfish numbers remained higher than target levels, the biomass of prey fishes decreased by a further 50%. While complete eradication of lionfish from the Caribbean is not likely, groups are actively removing them from coastal areas (mostly via spear and net). This study is a first step in showing that strategic local efforts that suppress the invasion to low levels can help protect and recover native fish communities affected by lionfish. This ground-breaking invasive lionfish research was conducted as a collaboration between REEF, Oregon State University, Simon Fraser University, and the Cape Eleuthera Institute.

Lotterhos, KE, SJ Dick, and DR Haggarty. 2014. Evaluation of rockfish conservation area networks in the United States and Canada relative to the dispersal distance for black rockfish (Sebastes melanops).

Evolutionary Applications. (2014) 238–259

REEF data were used to validate population estimates of Black Rockfish throughout western Canada, Washington State, and Oregon. These results were then used to evaluate the efficacy of marine reserve networks in these areas. The authors of the study estimated the scale of dispersal from genetic data in the black rockfish, and compared this estimate with the distance between Rockfish Conservation Areas that aim to protect this species (essentially evaluating whether the reserves are "connected" enough). Their findings showed that within each country, the distance between conservation areas was generally well connected. The distance between the networks in the two countries, however, was greater than the average dispersal per rockfish generation.

Jackson, AM, BX Semmens, Y Sadovy de Mitcheson, RS Nemeth, SA Heppell, PG Bush, A Aguilar-Perera, JAB Claydon, MC Calosso, KS Sealey, MT Schärer, G Bernardi. 2014. Population structure and phylogeography in Nassau Grouper (Epinephelus striatus), a mass-aggregating marine fish.

PLoS ONE. DOI: 10.1371/journal.pone.0097508

This study, co-authored by scientists from REEF's Grouper Moon Project, evaluated genetic connectedness between Nassau Grouper populations throughout the Caribbean. The authors obtained genetic tissue samples from 620 Nassau Grouper from 19 sites across 9 countries, including the Cayman Islands. They found evidence for strong genetic differentiation among Nassau Grouper subpopulations throughout the Caribbean. These results suggest that, despite a lack of physical barriers, Nassau grouper form multiple distinct subpopulations in the Caribbean Sea. Oceanography (regional currents, eddies) likely plays an important role in retaining larvae close to spawning sites at both local and regional spatial scales. These findings highlight the importance of conservation initiatives such at REEF's Grouper Moon program in the Cayman Islands.

Côté​, IM, L Akins, E Underwood, J Curtis-Quick, SJ Green. 2014. Setting the record straight on invasive lionfish control: Culling works.

PeerJ PrePrints. 2:e398v1

Frequent culling of the invasive Indo-Pacific Lionfish throughout the Caribbean has been shown to cause a shift towards more wary and reclusive behaviour by lionfish, which has prompted calls for halting culls. This paper, co-authored by REEF Lionfish Program researchers, addresses those concerns and reviews research conducted by REEF and other research efforts. Culling successfully lowers lionfish numbers and has been shown to stabilise or even reverse declines in native prey fish. Partial culling is often as effective as complete local eradication, yet requires significantly less time and effort. Abandoning culling altogether would therefore be seriously misguided and a hindrance to conservation. The authors also offer suggestions for how to design removal programs that minimize behavioural changes and maximize culling success.

Archer, SK, JE Allgeier, BX Semmens, SA Heppell, CV Pattengill-Semmens, AD Rosemond, PG Bush, CM McCoy, BC Johnson, CA Layman. 2014. Hot Moments in Spawning Aggregations: implications for ecosystem-scale nutrient cycling.

Coral Reefs. 10.1007/s00338-014-1208-4

This paper presents results from a study conducted as part of REEF's Grouper Moon Project, evaluating the potential ecosystem-level effect of Nassau Grouper aggregations. In particular, the study looked at the impact the spawning aggregation has in creating biogeochemical "hot moments", which occur when a temporary increase in availability of one or more limiting nutrients results in elevated rates of biogeochemical reactions. In this case, the limited nutrients are nitrogen and phosphorus, and the temporary increase is resulting from all of the grouper excrement that results when approximately 5,000 Nassau Grouper gather in a small area for 10 days during the spawning season.

Scyphers, SB, SP Powers, JL Akins, JM Drymon, CW Martin, ZH Schobernd, PJ Schofield, RL Shipp, and TS Switzer. 2014. The Role of Citizens in Detecting and Responding to a Rapid Marine Invasion.

Conservation Letters. doi: 10.1111/conl.12127

The authors of this study (including REEF staff Lad Akins) examined the recent case of Indo-Pacific lionfish invading Northern Gulf of Mexico coastal waters. The authors compared traditional reef fish monitoring efforts to less traditional data including the observations of divers through REEF's Volunteer Fish Survey Project and spearfishers. They found that citizen observations documented lionfish 1-2 years earlier and more frequently than the more traditional monitoring efforts. The authors also explored the willingness of spearfishers to help minimize impacts of the invasion by harvesting lionfish. They found that spearfishers who had encountered more lionfish while diving perceived them as more harmful to the habitat than less experienced divers and were also more likely to participate in harvesting initiatives. The authors also report that encouragement from scientists and managers was a far better motivator than the desire to harvest lionfish for personal consumption. This study demonstrates the value of engaging citizens for assessing and responding to large-scale and time-sensitive conservation problems.

Thorson, JT, MD Scheuerell, BX Semmens, and CV Pattengill-Semmens. 2014. Demographic Modeling of Citizen Science Data Informs Habitat Preferences and Population Dynamics of Recovering Fishes.


Managing natural populations and communities requires detailed information regarding demographic processes (or status of a population) at large spatial and temporal scales. This combination is challenging for both traditional scientific surveys, which often operate at localized scales, and citizen science designs, which often provide data with few auxiliary information (i.e. no information about individual age or condition). The authors of this study combine citizen science data collected at large scales (REEF Volunteer Fish Survey Project data) with recently developed statistical demographic modeling techniques. The model analysis included two managed reef fishes in the Gulf of Mexico to estimate demographic trends, habitat associations, and interannual variability in recruitment of Goliath Grouper and Mutton Snapper. The results identify strong preferences for artificial structure for the recovering Goliath Grouper, while revealing little evidence of either habitat associations or trends in abundance for Mutton Snapper. Results are also contrasted with a typical modeling approach to demonstrate the importance of accounting for the statistical complexities implied by spatially structured citizen science data. Results also highlight the utility and management benefits of combining demographic models and citizen science data.

Akins, JL, JA Morris Jr, and SJ Green. 2014. In Situ Tagging Technique for Fishes Provides Insight Into Growth and Movement of Invasive Lionfish.

Ecology and Evolution. 10.1002/ece3.1171

Information on fish movement and growth is primarily obtained through the marking and tracking of individuals with external tags, which are usually affixed to anesthetized individuals at the surface. However, the quantity and quality of data obtained by this method is often limited by small sample sizes owing to the time associated with the tagging process, high rates of tagging-related mortality, and displacement of tagged individuals from the initial capture location. To address these issues, we describe a technique for applying external streamer and dart tags in situ, which uses SCUBA divers to capture and tag individual fish on the sea floor without the use of anesthetic. We demonstrate this method for Indo-Pacific lionfish (Pterois volitans/P. miles), species which are particularly vulnerable to barotrauma when transported to and handled at the surface. To test our method, we tagged 161 individuals inhabiting 26 coral reef locations in the Bahamas over a period of 3 years. Our method resulted in no instances of barotrauma, reduced handling and recovery time, and minimal post-tagging release displacement compared with conventional ex situ tag application. Opportunistic resighting and recapture of tagged individuals reveals that lionfish exhibit highly variable site fidelity, movement patterns, and growth rates on invaded coral reef habitats. In total, 24% of lionfish were resighted between 29 and 188 days after tagging. Of these, 90% were located at the site of capture, while the remaining individuals were resighted between 200 m and 1.1 km from initial site of capture over 29 days later. In situ growth rates ranged between 0.1 and 0.6 mm/day. While individuals tagged with streamer tags posted slower growth rates with increasing size, as expected, there was no relationship between growth rate and fish size for individuals marked with dart tags, potentially because of large effects of tag presence on the activities of small bodied lionfish (i.e., <150 mm), where the tag was up to 7.6% of the lionfish's mass. Our study offers a novel in situ tagging technique that can be used to provide critical information on fish site fidelity, movement patterns, and growth in cases where ex situ tagging is not feasible.

Côté, IM, SJ Green, JA Morris Jr, JL Akins, D Steinke. 2013. Diet richness of invasive Indo-Pacific lionfish revealed by DNA barcoding.

Marine Ecology Progress Series. 72: 249–256

Invasive lionfish in the western Atlantic are known to be voracious predators. Their unusual hunting behavior suggests that they could prey on most fish species within their gape size limits. Significant research by REEF researchers and others has been conducted looking at stomach contents of lionfish to identify prey. However, relatively few prey species have been identified because of the challenge of identifying partly digested prey. It is also difficult to know how well the identifiable diet reflects the unidentified portion. The authors of this study addressed this issue by DNA-barcoding unidentifiable fish items from the stomachs of 130 lionfish. They identified 37 prey species, half of which had previously not been recorded. The visually identifiable species only accounted for 25% of the total prey items, making it clear that extrapolating total prey from the identifiable portion is not accurate. The barcoding technique used can increase the ability to predict the impacts of invasive predators on recipient communities.

Holt, BG, R Rioja-Nieto, MA MacNeil, J Lupton, and C Rahbek. 2013. Comparing Diversity Data Collected Using a Protocol Designed for Volunteers with Results from a Professional Alternative.

Methods in Ecology and Evolution. doi: 10.1111/2041-210X.12031

Research conducted by Dr. Ben Holt from University of East Angila in the UK shows that methods to record marine diversity used by REEF surveyors returned results consistent with techniques favoured by peer-reviewed science. The findings give weight to the growing phenomenon of citizen science programs such as REEF's Volunteer Survey Project. The field study compared methods used by REEF volunteer SCUBA divers with those used by professional scientists to measure the variety of fish species in three Caribbean sites in the Turks and Caicos. The divers surveyed the sites using two methods – the 'belt transect', used in peer reviewed fish diversity studies, and the 'roving diver technique', used by REEF volunteers. Two teams of 12 divers made 144 separate underwater surveys across the sites over four weeks. While the traditional scientific survey revealed sightings of 106 different types of fish, the volunteer technique detected greater marine diversity with a total of 137 in the same waters. Dr Holt led the research in partnership with the Centre for Marine Resource Studies in the Caribbean and the University of Copenhagen, Denmark. He said: "The results of this study are important for the future of citizen science and the use of data collected by these programs. Allowing volunteers to use flexible and less standardised methods has important consequences for the long term success of citizen science programs. Amateur enthusiasts typically do not have the resources or training to use professional methodology. Our study demonstrates the quality of data collected using a volunteer method can match, and in some respects exceed, protocols used by professional scientists."

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