FISH AGGREGATIONS AND BIOGEOCHEMICAL HOT SPOTS ACROSS REGIONAL ENVIRONMENTAL GRADIENTS

• Sponsoring Institution: National Institute of Food and Agriculture
• Project Status: COMPLETE
• Funding Source: HATCH
• Reporting Frequency: Annual
• Accession No.: 1001040
• Project Start Date: Sep 1, 2013
• Project End Date: Sep 30, 2017
• Program Code: [(N/A)]- (N/A)

Recipient Organization
NORTH CAROLINA STATE UNIV
(N/A)
RALEIGH,NC 27695

Performing Department
Biology

Non Technical Summary
Consumers have long been acknowledged for their role in top-down regulation of ecosystems (resource consumption), but their influence through bottom-up mechanistic pathways (resource supply) is often underappreciated. Namely, through physiological processes of consumption, assimilation and regeneration, consumers cycle nutrients that are readily available for biological uptake by primary producers and microorganisms. Despite seminal research over three decades ago, consumermediated nutrient supply has not been integrated into models of coastal ecosystem dynamics. Yet nutrient supply from consumers may be a regulating factor of ecosystem processes, especially when consumers aggregate, e.g., fishes around structurally complex habitat. Examining this bottom-up mechanistic pathway is essential for a more holistic understanding of seagrass ecosystems - one of the most important, yet threatened, ecosystems globally. Questions that frame the research include: (1) Does concentrated nutrient supply from consumers result in distinct biogeochemical hot spots in seagrass beds?, and (2) How do consumer effects on ecosystem processes vary across regional environmental contexts, i.e., across gradients in nutrient availability and fishing pressure? The experimental design incorporates environmental variation by selecting multiple sites within three biogeographic regions in the Caribbean (The Bahamas, Hispaniola, and Grenada/St.Vincent/Grenadines). The experiments will utilize artificial reefs that mimic natural patch reef habitats by serving to concentrate animals at high densities. Response variables reflecting ecosystem processes (e.g., seagrass nutrient content, seagrass biomass, primary producer diversity) will be measured at reef sites and compared with control sites (seagrass sites without reefs). The spatial extent over which ecosystem processes may be affected, i.e., distance from artificial reef, will be quantified and used to detect ecological thresholds in ecosystem responses. Predictor variables, including measures of ambient nutrient availability, fish densities and fish grazing rates, will be used to contextualize the relative importance of consumer-mediated nutrient supply. The hierarchical experimental design and two-pronged analysis will characterize relationships across environmental gradients (e.g., terrestrial land use practices) found among and within the biogeographic regions, facilitating a conceptual framework needed to predict when, where, and why consumer-mediated nutrient supply is an important control of ecosystems processes in seagrass beds. The project is designed to maximize broader impacts at regional, national, and local levels, through extensive integration of research and education efforts. The entire program is designed in partnership with The Nature Conservancy, for which the PI serves on the Caribbean Advisory Board. The Nature Conservancy is at the forefront of the Caribbean Challenge, an effort to protect 20% of marine habitats by the year 2020. All of the planned study sites are part of current, or proposed, marine protected areas as part of the Caribbean Challenge initiative. Data from the research will inform monitoring plans, management strategies and prioritization of conservation measures. For example, the site in Haiti will be within boundaries of the country's first proposed marine protected area, the design of which the PI is working with The Nature Conservancy to designate. Outreach initiatives at local and national levels, will be designed with sensitivity to local environmental and social opportunities/challenges. We will work with different local partners in each country, crafting educational efforts that are best suited to challenges and opportunities at each individual site. For example, in The Bahamas, the PI will help design and develop a new center on Abaco Island intended to support research, education and conservation activities: ACRE (Abaco Conservation, Research & Education Center). The outreach website The Abaco Scientist will also be expanded, continuing to provide a platform for quick dissemination of information to the general public. Emphasis will be placed on training minority students underrepresented in the sciences through FIU, providing experiences within an integrated science, education, and cross-cultural integration model. As such, the project will provide for broad advances regarding the function of seagrass ecosystems, while being embedded in an extensive outreach context in which conservation and education benefits will be prioritized.

Animal Health Component

(N/A)

Research Effort Categories

Basic

100%

Applied

(N/A)

Developmental

(N/A)

Classification

Knowledge Area (KA)	Subject of Investigation (SOI)	Field of Science (FOS)	Percent
135	0330	1070	100%

Knowledge Area
135 - Aquatic and Terrestrial Wildlife;

Subject Of Investigation
0330 - Wetland and riparian systems;

Field Of Science
1070 - Ecology;

Keywords

land use practices

eutrophication

food web

fish

estuary

runoff

Goals / Objectives
An underappreciated aspect of the functioning of coastal ecosystems is the supply of nutrients by fishes to aquatic primary producers. The importance of this nutrient pathway like varies as a function of the background environmental conditions (e.g., ambient nutrient levels are higher in water bodied adjacent to agricultural areas). The goal of this project is examine how different landuse practices affects the role of fishes as nutrient vectors.

Project Methods
The experimental units for this project are artificial reefs constructed with cinder blocks in seagrass beds. We will monitor various aspects of the seagrass beds including seagrass growth rate and nutrient content. We also will sample ambient nutrients at each site providing information on ambient site conditions, that then can belinked back to terrestrial land use practices.

Progress 10/01/16 to 09/30/17

Outputs
Target Audience:The scientific community and general public. Changes/Problems:None What opportunities for training and professional development has the project provided?Working with dozens of students and community members. How have the results been disseminated to communities of interest?https://appliedecology.cals.ncsu.edu/absci/ craiglaymanlab.com What do you plan to do during the next reporting period to accomplish the goals?Continue the research outlined in the grant.

Impacts
What was accomplished under these goals? Understanding these dynamics better.

Publications

• Type: Journal Articles Status: Accepted Year Published: 2017 Citation: Archer, S.K., Stevens, J.L., Rossi, R.E., Matterson, K. and Layman, C.A. 2017. Abiotic conditions drive significant variability in nutrient processing by a common Caribbean sponge, Ircinia felix. Limnology and Oceanography.
• Type: Journal Articles Status: Published Year Published: 2017 Citation: Sweatman, J.L., Layman, C.A., and Fourqurean, J.W. 2017. Habitat fragmentation has weak impacts on aspects of ecosystem functioning in a sub-tropical seagrass bed. Marine Environmental Research 126:95-108.
• Type: Journal Articles Status: Published Year Published: 2017 Citation: Allgeier, J.E., Burkepile, D.E., and Layman, C.A. 2017. Animal pee in the sea: consumer- mediated nutrient dynamics in the worlds oceans. Global Change Biology. doi: 10.1111/gcb.13625.
• Type: Journal Articles Status: Published Year Published: 2016 Citation: Layman, C.A., Allgeier, J.E., and Monta�a, C.G. 2016. Mechanistic evidence of enhanced production on artificial reefs: A case study in a Bahamian seagrass ecosystem. Ecological Engineering. 95:574-579.
• Type: Journal Articles Status: Published Year Published: 2016 Citation: Allgeier, J.E, Valdivia, A., Cox, C., Bruno, J.F., and Layman, C.A. 2016. Fishing down nutrients on coral reefs. Nature Communications DOI: 10.1038/ncomms12461.
• Type: Journal Articles Status: Published Year Published: 2016 Citation: Stoner, E.W., Sebilian, S.S., Layman, C.A. 2016. Zooxanthellae densities in upside-down jellyfish, Cassiopea spp., from coastal habitats of The Bahamas. Revista de Biolog�a Marina y Oceanograf�a. 51:203-208.

Progress 10/01/15 to 09/30/16

Outputs
Target Audience:Any stakeholder in theworld's oceans. Changes/Problems:None What opportunities for training and professional development has the project provided?It has provided numerous aopportunities for undergraduates, e.g., almost 3,000 hours in 2015 alone:http://craiglaymanlab.com/wp-content/uploads/Layman-Lab-2015-Undergrad-Hours-and-Stats.pdf How have the results been disseminated to communities of interest?Yes - through the publication process and these websites: https://appliedecology.cals.ncsu.edu/absci/ http://craiglaymanlab.com/ What do you plan to do during the next reporting period to accomplish the goals?Continue the protocols outlined in the grant proposal.

Impacts
What was accomplished under these goals? We have continued to monitor the core experimental units (artifically reefs) annually, with periodic side publications along the way.

Publications

• Type: Journal Articles Status: Awaiting Publication Year Published: 2017 Citation: Allgeier, J.E., Burkepile, D.E., and Layman, C.A. In press. Animal pee in the sea: consumer- mediated nutrient dynamics in the worlds oceans. Global Change Biology.
• Type: Journal Articles Status: Accepted Year Published: 2016 Citation: Layman, C.A., Allgeier, J.E., and Monta�a, C.G. 2016. The attraction-production debate viewed from the bottom-up: mechanistic evidence of increased production thresholds. Ecological Engineering. 95:574-579.
• Type: Journal Articles Status: Published Year Published: 2016 Citation: Allgeier, J.E, Valdivia, A., Cox, C., Bruno, J.F., and Layman, C.A. 2016. Fishing down nutrients on coral reefs. Nature Communications DOI: 10.1038/ncomms12461.
• Type: Journal Articles Status: Published Year Published: 2016 Citation: Stoner, E.W., Sebilian, S.S., Layman, C.A. 2016. Zooxanthellae densities in upside-down jellyfish, Cassiopea spp., from coastal habitats of The Bahamas. Revista de Biolog�a Marina y Oceanograf�a. 51:203-208.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Layman, C.A., Giery, S.T., Buhler, S., Rossi, R., Penland, T., Henson, M.N., Bogdanoff, A.K., Cove, M.V., Irizarry, A.D., Schalke, C.M., and Archer, S.K. 2015. A primer on the history of food web ecology: Fundamental contributions of fourteen researchers. Food Webs 4:14-24.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Allgeier, J.E., Wegner, S., Rosemond, A.D., Schindler, D.E., and Layman, C.A. 2015. Metabolic theory and biodiversity, but not stoichiometry, best predict nutrient recycling in a diverse food web. Proceedings of the National Academy of Sciences 112(20):E2640-E2647.

Progress 10/01/14 to 09/30/15

Outputs
Target Audience:International scientists for the research component. The results have direct implications for citizens of The Bahamas and Haiti, but more broadly citizens of the US and any other country that have seagrass ecosystems. Changes/Problems:None What opportunities for training and professional development has the project provided?The project helped support almost 3,000 of research activities for NCSU undergraduates:http://craiglaymanlab.com/wp-content/uploads/Layman-Lab-2015-Undergrad-Hours-and-Stats.pdf. It also supported the research of 3 graduate students. I base much of my course (Applied Ecology) around research conducted with this funding. It also contributed to the development of environmental NGOs in The Bahamas. How have the results been disseminated to communities of interest?Scientific publications, our outreach website (http://appliedecology.cals.ncsu.edu/absci/) and myriad outreach activities (http://craiglaymanlab.com/outreach/). What do you plan to do during the next reporting period to accomplish the goals?We plan to continue to extend and expand all activities mentioned herein.

Impacts
What was accomplished under these goals? The publications listed represent the many ways we have progressed in this area. We continue to compile data on how environmenal context mediates the role of fishes in coastal ecosystems.

Publications

• Type: Journal Articles Status: Published Year Published: 2015 Citation: Fodrie, J.F., Yeager, L.A., Grabowski, J.H., Layman, C.A., Sherwood, G.D., and Kenworthy, M.D. 2015. Measuring individuality in habitat use across complex landscapes: approaches, constraints, and implications for assessing resource specialization. Oecologia 178(1):75-87.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Layman, C.A., Newsome, S.D., and Crawford, T.G. 2015. Individual specialization within populations: emerging areas of study. Oecologia 178(1):1-4.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Archer, S.K., Stoner, E.W., and Layman, C.A. 2015. Context dependent species interaction between a sponge (Halichondria melanadocia) and a seagrass (Thalassia testudinum). Journal of Marine Biology and Ecology 465:33-40.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Stoner, E.W. and Layman, C.A. 2015. Bristle worms attack: Benthic jellyfish are not trophic dead ends. Frontiers in Ecology and the Environment 13:226-227.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Allgeier, J.E., Wegner, S., Rosemond, A.D., Schindler, D.E., and Layman, C.A. 2015. Metabolic theory and biodiversity, but not stoichiometry, best predict nutrient recycling in a diverse food web. Proceedings of the National Academy of Sciences 112(20):E2640-E2647.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Giery, S.T., Layman, C.A. and Langerhans, R.B. 2015. Anthropogenic ecosystem fragmentation drives shared and unique patterns of sexual signal divergence among three species of Bahamian mosquitofish. Evolutionary Applications 8(7):679-691.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Layman, C.A., Giery, S.T., Buhler, S., Rossi, R., Penland, T., Henson, M.N., Bogdanoff, A.K., Cove, M.V., Irizarry, A.D., Schalke, C.M., and Archer, S.K. 2015. A primer on the history of food web ecology: Fundamental contributions of fourteen researchers. Food Webs 4:14-24.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Riesch, R., Easter, T., Layman, C.A., and Langerhans, R.B. 2015. Rapid human-induced divergence of life history strategies in Bahamian live-bearing fishes (family Poeciliidae). Journal of Animal Ecology 84(6):1732-1743.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Heinen-Kay, J.L., Noel, H.G., Layman, C.A., and Langerhans, R.B. 2014. Human-caused habitat fragmentation drives rapid divergence of male genital morphology in Bahamian mosquitofish. Evolutionary Applications 7(10):1252-1267.

Progress 10/01/13 to 09/30/14

Outputs
Target Audience: The scientific community, as well as citizens of Haiti and The Bahamas Changes/Problems: Research was so successful in Haiti, we have shifted from 3 sites in each of Curacao, The Bahamas, Haiti, and Grenada (12 total), to 23 sites across 4 islands in Haiti and The Bahamas. This will give our experimental design much more statistical power. A significant improvment in design. What opportunities for training and professional development has the project provided? I have leveraged funds to help train 2 post-docs, 2 graduate students, 6 undergraduates and dozens of local community members in the countries where research is based. How have the results been disseminated to communities of interest? More than 10 community meetings and this website: http://appliedecology.cals.ncsu.edu/absci/ What do you plan to do during the next reporting period to accomplish the goals? Follow the outlined timeline in the funded grant proposal.

Impacts
What was accomplished under these goals? In the first year of the project, we are still accumulating data. But the research has already lead to establishment of the first marine protected area ever in Haiti. Multiple publications resulted from the first year of research.

Publications

• Type: Journal Articles Status: Accepted Year Published: 2015 Citation: Layman, C.A., Newsome, S.D., and Crawford, T.G. In press. Individual specialization within populations: emerging areas of study. Oecologia.
• Type: Journal Articles Status: Accepted Year Published: 2014 Citation: Zapata, M.J., Yeager, L.A., and Layman, C.A. 2014. Day-night patterns in natural and artificial patch reef fish assemblages of The Bahamas. Caribbean Naturalist 18:1-15.
• Type: Journal Articles Status: Accepted Year Published: 2015 Citation: Allgeier, J.E., Layman, C.A., Mumby, P.J., and Rosemond, A.D. In press. Biogeochemical implications of biodiversity and community structure across multiple coastal ecosystems. Ecological Monographs.
• Type: Journal Articles Status: Accepted Year Published: 2015 Citation: Archer, S.K., Allgeier, J.E., Semmens, B.X., Heppell, S.A., Pattengill-Semmens, C.V., Rosemond, A.D., Bush, P.G., McCoy, C.M., Johnson, B.C., and Layman, C.A. In press. Hot moments in spawning aggregations: implications for ecosystem-scale nutrient cycling. Coral Reefs DOI 10.1007/s00338-014-1208-4.
• Type: Journal Articles Status: Accepted Year Published: 2014 Citation: Yeager, L.A., Stoner, E.W., Zapata, M.J., and Layman, C.A. 2014. Does landscape context mediate the nature of density dependence for a coral reef fish? Ecological Applications 24:1833-1841.
• Type: Journal Articles Status: Accepted Year Published: 2013 Citation: Burkepile, D.E., Allgeier, J.A., Shantz, A.A., Pritchard, C.E., Lemoine, N., Bhatti, L., and Layman, C.A. 2013. Nutrient supply from fishes facilitates macroalgae and suppresses corals in a Caribbean coral reef ecosystem. Scientific Reports 3:1493 DOI 10.1038.
• Type: Journal Articles Status: Accepted Year Published: 2014 Citation: Allgeier, J.E., Layman, C.A., Mumby, P.J., Rosemond, A.D. 2014. Consistent nutrient storage and supply mediated by diverse fish communities in coral reef ecosystems. Global Change Biology 20(8):2459-2472.

Progress 09/01/13 to 09/30/13

Outputs
Target Audience: Both the broader scientific community and local communities and stakeholders in The Bahamas and Haiti. Changes/Problems: Research was so successful in Haiti, we have shifted from3 sites in each of Curacao, The Bahamas, Haiti, and Grenada (12 total), to 23 sites across 4 islands in Haiti and The Bahamas. This will give our experimental design much more statistical power. A significant improvment in design. What opportunities for training and professional development has the project provided? I have leveraged funds to help train 2 post-docs, 2 graduate students, 6 undergraduates and dozens of local community members in the countries where research is based. How have the results been disseminated to communities of interest? More than 10 community meetings and this website: http://appliedecology.cals.ncsu.edu/absci/ What do you plan to do during the next reporting period to accomplish the goals? Follow the outlined timeline in the funded grant proposal.

Impacts
What was accomplished under these goals? In the first year of the project, we are still accumulating data. But the research has already lead to establishment of the first marine protected area ever in Haiti. Multiple publications resulted from the first year of research.

Publications

• Type: Journal Articles Status: Published Year Published: 2014 Citation: Zapata, M.J., Yeager, L.A., and Layman, C.A. 2014. Day-night patterns in natural and artificial patch reef fish assemblages of The Bahamas. Caribbean Naturalist 18:1-15.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Allgeier, J.E., Layman, C.A., Mumby, P.J., and Rosemond, A.D. In press. Biogeochemical implications of biodiversity and community structure across multiple coastal ecosystems. Ecological Monographs.
• Type: Journal Articles Status: Published Year Published: 2015 Citation: Archer, S.K., Allgeier, J.E., Semmens, B.X., Heppell, S.A., Pattengill-Semmens, C.V., Rosemond, A.D., Bush, P.G., McCoy, C.M., Johnson, B.C., and Layman, C.A. In press. Hot moments in spawning aggregations: implications for ecosystem-scale nutrient cycling. Coral Reefs DOI 10.1007/s00338-014-1208-4.
• Type: Journal Articles Status: Published Year Published: 2014 Citation: Yeager, L.A., Stoner, E.W., Zapata, M.J., and Layman, C.A. 2014. Does landscape context mediate the nature of density dependence for a coral reef fish? Ecological Applications 24:1833-1841.
• Type: Journal Articles Status: Published Year Published: 2014 Citation: Allgeier, J.E., Layman, C.A., Mumby, P.J., Rosemond, A.D. 2014. Consistent nutrient storage and supply mediated by diverse fish communities in coral reef ecosystems. Global Change Biology 20(8):2459-2472.
• Type: Journal Articles Status: Published Year Published: 2013 Citation: Burkepile, D.E., Allgeier, J.A., Shantz, A.A., Pritchard, C.E., Lemoine, N., Bhatti, L., and Layman, C.A. 2013. Nutrient supply from fishes facilitates macroalgae and suppresses corals in a Caribbean coral reef ecosystem. Scientific Reports 3:1493 DOI 10.1038.
• Type: Journal Articles Status: Accepted Year Published: 2015 Citation: Layman, C.A., Newsome, S.D., and Crawford, T.G. In press. Individual specialization within populations: emerging areas of study. Oecologia.