Description: Author Posting. © National Research Council Canada, 2004. This article is posted here by permission of National Research Council Canada for personal use, not for redistribution. The definitive version was published in Canadian Journal of Fisheries and Aquatic Sciences 61 (2004): 1954-1964, doi:10.1139/F04-126.

Description: In many long-lived species such as Arctic grayling (Thymallus arcticus), population growth rate is most sensitive to changes in adult survival probabilities. Understanding the factors that regulate adult survival in this species should provide insight into the population dynamics of this and other long-lived Arctic species. Using the program MARK, we analyzed 17 years of mark–recapture data to estimate survival rates for Arctic grayling in the Kuparuk River, Alaska, from 1985 to 2000. Mean annual survival rates overall ranged from 0.39 to 1.0 and averaged 0.71 ± 0.05 for resident and 0.75 ± 0.05 for nonresident fish. Spending the summer in the more productive fertilized zone of the experimental reach had no influence on survival despite higher productivity on all trophic levels and consistently higher growth rates in Arctic grayling. None of the environmental (stream temperature, discharge, winter severity, and incidence of drought) or population parameters (growth, condition factor, and mean fish size) that we examined explained significant amounts of variance in survival rates. The lack of responsiveness of survival to annual environmental conditions was unexpected and suggests that multiyear factors or life history tactics that maintain survival at the expense of growth and fecundity likely determine survival.

Description: This research was supported by National Science Foundation grants OPP-9911278 and DEB-9810222 in conjunction with the Long-term Ecological Research Program.

Description: Author Posting. © The Author(s), 2007. This is the author's version of the work. It is posted here by permission of Ecological Society of America for personal use, not for redistribution. The definitive version was published in Ecological Applications 17, Suppl. (2007): S42–S63, doi:10.1890/06-0452.1.

Description: The sustainability of coastal ecosystems in the face of widespread environmental change is an issue of pressing concern throughout the world (Emeis et al. 2001). Coastal ecosystems form a dynamic interface between terrestrial and oceanic systems and are one of the most productive ecosystems in the world. Coastal systems probably serve more human uses than any other ecosystem and they have always been valued for their rich bounty of fish and shellfish. Coastal areas are also the sites of the nation’s and the world’s most intense commercial activity and population growth; worldwide, approximately 75% of the human population now lives in coastal regions (Emeis et al. 2001). Over the past three decades nutrient enrichment of coastal and estuarine waters has become the premier issue for both scientists and managers (National Research Council 2000). Our understanding of coastal eutrophication has been developed principally through monitoring of estuaries, with a focus on pelagic or subtidal habitats (National Research Council 2000, Cloern 2001). Because estuarine systems are usually nitrogen limited, NO3- is the most common nutrient responsible for cultural nutrient enrichment (Cloern 2001). Increased nitrogen delivery to pelagic habitats of estuaries produces the classic response of ecosystems to stress (altered primary producers and nutrient cycles and loss of secondary producer species and production; Nixon 1995, Rapport and Whitford 1999, Deegan et al. 2002). Salt marsh ecosystems have been thought of as not susceptible to nitrogen over-loading because early studies found added nitrogen increased marsh grass production (primarily Spartina spp., cordgrass) and concluded that salt marshes can adsorb excess nutrients in plants and salt marsh plant-derived organic matter as peat (Verhoeven et al. 2006). Detritus from Spartina is important in food webs (Deegan et al. 2000) and in creating peat that forms the physical structure of the marsh platform (Freidrichs and Perry 2001). However, the accumulation of peat and inputs of sediments and loss of peat through decomposition and sediment through erosion may be altered under high nutrient regimes and threaten the long-term stability of marsh systems. Nitrogen addition may lead to either net gain or loss of the marsh depending on the balance between increased marsh plant production and increased decomposition. Absolute change in marsh surface elevation is determined by marsh plant species composition, production and allocation to above- and belowground biomass, microbial decomposition, sedimentation, erosion and compaction (Friedrichs and Perry 2001). Levine et al. (1998) suggested that competitive dynamics among plants might be affected by nutrient enrichment, potentially altering relative abundance patterns favoring species with less belowground storage and thus lowering rates of peat formation. When combined with the observation that nutrient additions may also stimulate microbial respiration and decomposition (Morris and Bradley 1999), the net effect on the salt marsh under conditions of chronic nitrogen loading is a critical unknown. Although most research treats nutrient enrichment as a stand-alone stress, it never occurs in isolation from other perturbations. The effect of nutrient loading on species composition (both plants and animals) and the resultant structure and function of wetlands has been largely ignored when considering their ability to adsorb nutrients (Verhoeven et al. 2006). Recent studies suggest the response of estuaries to stress may depend on animal species composition (Silliman et al. 2005). Animal species composition may alter the balance between marsh gain and loss as animals may increase or decrease primary production, decomposition or N recycling (Pennings and Bertness 2001). Failure to understand interactions between nutrient loading and change in species composition may lead to underestimating the impacts of these stresses. The 'bottom up or top down' theory originated from the observation that nutrient availability (bottom up)sets the quantity of primary productivity, while other studies have shown that species composition (top down), particularly of top consumers, has a marked and cascading effect on ecosystems, including controlling species composition and nutrient cycling (Matson and Price 1992, Pace et al. 1999). Most examples of trophic cascades are in aquatic ecosystems with fairly simple, algal grazing pelagic food webs (Strong 1992). The rarity of trophic cascades in terrestrial systems has been attributed to the importance of detrital food webs (Polis 1999). Detritus-based aquatic ecosystems, such as salt marshes, bogs, and swamps, have classically been considered bottom-up or physically controlled ecosystems. Recent experiments, however, suggest that salt marshes may exhibit top-down control at several trophic levels (Silliman and Zeiman. 2001, Silliman and Bertness 2002, Quiñones-Rivera and Fleeger 2005). One abundant, ubiquitous predator, a small (〈10 cm total length) killifish (Fundulus heteroclitus, mummichog) has been suggested to control benthic algal through a trophic cascade because they prey on the invertebrates that graze on the benthic algae (Kneib 1997, Sarda et al. 1998). In late summer, killifish are capable of consuming 3-10 times the creek meiofauna production and meiofauna in the absence of predators appear capable of grazing over 60% of the microalgal community per day (Carman et al. 1997). Strong top-down control by grazers is considered a moderating influence on the negative effects of elevated nutrients on algae (Worm et al. 2000). Small-scale nutrient additions and predator community exclusion experiments have demonstrated bottom-up and top-down control of macroinfauna in mudflats associated with salt marsh creeks (Posey et al. 1999, Posey et al. 2002). Together, these observations suggest mummichogs are at the top of a trophic cascade that controls benthic algae (Sarda et al. 1998). Mummichogs are also omnivorous and ingest algae, bulk detritus and the attached microbial community (D’Avanzo and Valiela 1990). As a result, marsh decomposition rates may be limited by top-down controls through trophic pathways or by release from competition with algae for nutrients. Whole-ecosystem experiments have shown that responses to stress are often not predictable from studies of the individual components (Schindler 1998). Developing the information needed to predict the interacting impacts of nutrient loading and species composition change requires experiments with realistic alterations carried out at scales of space and time that include the complexities of real ecosystems. Whole ecosystem manipulation experiments have been used effectively in other ecosystems (Bormann and Likens 1979, Carpenter et al. 1995), but they are rare in coastal research. Experiments in salt marshes have traditionally been less than a few m2. Our understanding of the response of salt marsh plants to nutrient enrichment is from small (〈10 m2), plot-level additions where uniform levels of dry inorganic fertilizer (20 to 〉 1000 g N m-2 y-1) are sprinkled on the marsh surface at low tide. Dry fertilizer additions were usually made every two weeks or monthly and the duration of elevated nutrient levels after these additions was usually not determined. Tidal water is the primary vector for N delivery to coastal marshes, suggesting that dry fertilizer addition to the marsh surface may not be the best basis for determining if Spartina production responds to nutrient enrichment of tidal waters. Similarly, our understanding of top-down controls in salt marshes also relies on small (1 - 4 m2) exclusion experiments that use cages to isolate communities from top consumers. While the design of these cage experiments has improved, there are some remaining drawbacks. For example, it is impossible to selectively exclude single species using cages, and recruitment or size-selective movement into or out of the cages may obscure interpretations. In addition, while these small-scale experiments provide insight into controls on isolated ecosystem processes, they do not allow for interaction among different parts of the ecosystem which may buffer or alter the impacts and are not appropriate for determining the effects of populations of larger more motile animals on whole-ecosystems or the effects of ecosystem changes on populations. For example, interactions may be caused when a motile species alters its distribution among the habitats available to it because of an experimental treatment. Small-scale experiments generally do not allow such events to happen. Complex feedbacks among physical and biological processes can alter accumulation rates and affect marsh elevation relative to sea level rise making extrapolation of small plot level experiments to whole marsh ecosystems problematic. We are conducting an ecosystem-scale, multi-year field experiment including both nutrient and biotic manipulations to coastal salt marsh ecosystems. We are testing, for the first time at the ecosystem level, the hypothesis that nutrient enrichment and species composition change have interactive effects across multiple levels of biological organization and a range of biogeochemical processes. We altered whole salt marsh creek watersheds (~60,000 m2 of saltmarsh) by addition of nutrients (15x ambient) in flooding waters and by a 60% reduction of a key fish species, the mummichog. Small marsh creek watersheds provide an ideal experimental setting because they have the spatial complexity, species composition and processes characteristic of the larger salt marsh ecosystem, which are often hundreds of thousands of m2. Manipulating entire salt marsh creeksheds allowed us to examine effects on large motile animals and the interactive effects of motile species changes on ecosystem processes without cage artifacts. Because our manipulations were done on whole-marsh ecosystems, we are able to evaluate the integrated and interactive effects on all habitats (e.g., water column, tidal creeks and marsh) and on populations. These experiments are similar in many respects to the small watershed experiments carried out in forested catchments. Our nutrient enrichment is novel compared to past studies in two important ways. We added nutrients (N and P) directly to the flooding tidal creek waters to mimic the way in which anthropogenic nutrients reach marsh ecosystems. All previous experimental salt marsh nutrient enrichment studies used a dose-response design with spatially uniform dry fertilizer loading on small plots (〈10 m2). Nutrients carried in water will interact and reach parts of the ecosystem differently than dry fertilizer. Our enrichment method also creates a spatial gradient of nutrient loading across the landscape that is proportional to the frequency and depth of inundation in the marsh. Spatial gradients in loading within an ecosystem are typical in real world situations in many terrestrial and aquatic ecosystems. Because of our enrichment method, at any location in the ecosystem, nutrient load will be a function of the nutrient concentration in the water, the frequency and depth of tidal flooding and the reduction of nutrients from the flooding waters by other parts of the ecosystem. Uniform loading misses important aspects of the spatial complexity of ecosystem exposure and response. This work is organized around two questions that are central to understanding the long-term fate of coastal marshes: 1. Does chronic nutrient enrichment via flooding water increase primary production more than it stimulates microbial decomposition? 2. Do top-down controls change the response of the salt marsh ecosystem to nutrient enrichment? Here we present findings on the first 2 years of these experiments including 1) water chemistry, 2) standing stocks and species composition of benthic microalgae, 3) microbial production, 4) species composition and ecophysiology of macrophytes, 5) invertebrates, and 6) nekton. Because even highly eutrophic waters result in nutrient loading that is an order of magnitude less than most plot level experiments, we expected little stimulation of salt marsh vascular plant growth. However, moderate levels of nutrient enrichment in the water column were expected to increase benthic algal biomass and to stimulate bacterial activity and detrital decomposition throughout the ecosystem because of direct uptake of nitrogen from the water column and availability of more high quality organic matter from increased algal production. We predicted nutrient enrichment would increase invertebrate production because of an increase of high quality microalgal and microbial production at the base of the food web. Finally, we predicted that fish reduction would reduce predation on benthic invertebrates resulting in increased abundance of benthic invertebrates that would graze down the benthic algae.

Description: The National Science Foundation (Grant DEB 0213767, OCE 9726921, and OCE 0423565) supported this work. Additional funding was provided by the National Science Foundation postdoctoral fellowship in Microbial Biology (DBI-0400819), the NOAA Coastal Intern grant (NA04NOS4780182), the Office of Environmental Education of Louisiana, Middlebury College and Connecticut College.

Description: Author Posting. © The Author(s), 2007. This is the author's version of the work. It is posted here by permission of Elsevier B.V. for personal use, not for redistribution. The definitive version was published in Journal of Experimental Marine Biology and Ecology 357 (2008): 20-34, doi:10.1016/j.jembe.2007.12.003.

Description: Responses of infaunal saltmarsh benthic invertebrates to whole-ecosystem fertilization and predator removal were quantified in Plum Island Estuary, Massachusetts, USA. Throughout a growing season, we enriched an experimental creek on each flooding tide to 70 mM NO3 - and 4 mM PO4 -3 (a 10 x increase in loading above background), and we reduced Fundulus heteroclitus density by 60% in a branch of the fertilized and a reference creek. Macroinfauna and meiofauna were sampled in creek (mudflat and creek wall), marsh edge (tall form Spartina alterniflora) and marsh platform (Spartina patens and stunted S. alterniflora) habitats before and after treatments were begun; responses were tested with BACI-design statistics. Treatment effects were most common in the mid-range of the inundation gradient. Most fertilization effects were on creek wall where ostracod abundance increased, indices of copepod reproduction increased and copepod and annelid communities were altered. These taxa may use epiphytes (that respond rapidly to fertilization) of filamentous algae as a food source. Killifish reduction effects on meiobenthic copepod abundance were detected at the marsh edge and suggest predator limitation. Fish reduction effects on annelids did not suggest top-down regulation in any habitat; however, fish reduction may have stimulated an increased predation rate on annelids by grass shrimp. Interactions between fertilization and fish reduction occurred under S. patens canopy where indirect predator reduction effects on annelids were indicated. No effects were observed in mudflat or stunted S. alterniflora habitats. Although the responses of infauna to fertilization and predator removal were largely independent and of similar mild intensity, our data suggests that the effects of ecological stressors vary across the marsh landscape.

Description: This research was supported by the National Science Foundation under Grants No. 0213767 and 9726921.

Description: Author Posting. © The Author(s), 2013. This is the author's version of the work. It is posted here by permission of Inter-Research for personal use, not for redistribution. The definitive version was published in Marine Ecology Progress Series 492 (2013): 211-222, doi:10.3354/meps10495.

Description: The effects of eutrophication on coastal plants and sessile animals are becoming well known, but responses of mobile species are less well studied. Here, we link variation in abundance, biomass, body size, growth rate, and resource utilization in mummichogs (Fundulus heteroclitus) 〉 40 mm in length to experimental nutrient enrichment in Plum Island Sound, Massachusetts, USA. To mimic cultural eutrophication, dissolved fertilizer was released into replicate saltmarsh creeks on each rising tide throughout entire growing seasons. In the summer of the 6th year of enrichment, we released coded-wire tagged mummichogs into nutrient-enriched (n = 3733 fish) and reference (n = 3894 fish) creeks and recaptured them over the next two months. We found increased abundance (by 37%), biomass (58%), body size (8%), and herbivory (115%, measured as photosynthetic gut pigment content) in nutrient-enriched creeks, although body condition was unaffected. However, individual growth rates were 43% lower in nutrient-enriched creeks. Nutrient enrichment stimulated primary production causing a bottom-up enrichment of the food web, which fostered increased biomass and body size. However, the reduction in growth rate indicates an adverse consequence of long-term nutrient enrichment. This negative effect occurred in the absence of increased hypoxia in these highly tidally (4-m amplitude) flushed study creeks. The mummichog is an important predator/grazer in salt marshes, and nutrient-induced alterations in biomass or resource utilization will directly or indirectly affect lower trophic levels, including benthic algae, thereby impacting the 63 ecosystem-wide response to eutrophication.

Description: This material is based upon the work supported by the National Science Foundation under Grant Nos. 0816963, 0923689 and 0423565.

Description: This paper is not subject to U.S. copyright. The definitive version was published in Fishery Bulletin 107 (2009): 329-338.

Description: For most migratory fish, little is known about the location and size of foraging areas or how long individuals remain in foraging areas, even though these attributes may affect their growth, survival, and impact on local prey. We tested whether striped bass (Morone saxatilis Walbaum), found in Massachusetts in summer, were migratory, how long they stayed in non-natal estuaries, whether observed spatial patterns differed from random model predictions, whether fish returned to the same area across multiple years, and whether fishing effort could explain recapture patterns. Anchor tags were attached to striped bass that were caught and released in Massachusetts in 1999 and 2000, and recaptured between 1999 and 2007. In fall, tagged striped bass were caught south of where they were released in summer, confirming that fish were coastal migrants. In the first summer, 77% and 100% of the recaptured fish in the Great Marsh and along the Massachusetts coast, respectively, were caught in the same place where they were released. About two thirds of all fish recaptured near where they were released were caught 2–7 years after tagging. Our study shows that smaller (400–500 mm total length) striped bass migrate hundreds of kilometers along the Atlantic Ocean coast, cease their mobile lifestyle in summer when they use a relatively localized area for foraging (〈20 km2), and return to these same foraging areas in subsequent years.

Description: This project was administered through the Massachusetts Cooperative Fish and Wildlife Research Unit. The Massachusetts Cooperative Fish and Wildlife Research Unit is an association among the U.S. Geological Survey; University of Massachusetts Department of Natural Resources Conservation; Massachusetts Division of Marine Fisheries; Massachusetts Division of Fisheries and Wildlife, and the Wildlife Management Institute.

Description: Author Posting. © The Author(s), 2010. This is the author's version of the work. It is posted here by permission of Nature Publishing Group for personal use, not for redistribution. The definitive version was published in The ISME Journal 5 (2011): 1540–1548, doi:10.1038/ismej.2011.22.

Description: Functional redundancy in bacterial communities is expected to allow microbial assemblages to survive perturbation by allowing continuity in function despite compositional changes in communities. Recent evidence suggests, however, that microbial communities change both composition and function as a result of disturbance. We present evidence for a third response: resistance. We examined microbial community response to perturbation caused by nutrient enrichment in salt marsh sediments using deep pyrosequencing of 16S rRNA and functional gene microarrays targeting the nirS gene. Composition of the microbial community, as demonstrated by both genes, was unaffected by significant variations in external nutrient supply, despite demonstrable and diverse nutrient–induced changes in many aspects of marsh ecology. The lack of response to external forcing demonstrates a remarkable uncoupling between microbial composition and ecosystem-level biogeochemical processes and suggests that sediment microbial communities are able to resist some forms of perturbation.

Description: Funding for this research came from NSF(DEB-0717155 to JEH, DBI-0400819 to JLB). Support for the sequencing facility came from NIH and NSF (NIH/NIEHS-P50-ES012742-01 and NSF/OCE 0430724-J Stegeman PI to HGM and MLS, and WM Keck Foundation to MLS). Salary support provided from Princeton University Council on Science and Technology to JLB. Support for development of the functional gene microarray provided by NSF/OCE99-081482 to BBW. The Plum Island fertilization experiment was funded by NSF (DEB 0213767 and DEB 0816963).

Description: Author Posting. © The Oceanography Society, 2013. This article is posted here by permission of The Oceanography Society for personal use, not for redistribution. The definitive version was published in Oceanography 26, no. 3 (2013): 168–179, doi:10.5670/oceanog.2013.60.

Description: The paucity of data on migratory connections and an incomplete understanding of how mobile organisms use geographically separate areas have been obstacles to understanding coastal dynamics. Research on acoustically tagged striped bass (Morone saxatilis) at the Plum Island Ecosystems (PIE) Long Term Ecological Research site, Massachusetts, documents intriguing patterns of biotic connectivity (i.e., long-distance migration between geographically distinct areas). First, the striped bass tagged at PIE migrated southward along the coast using different routes. Second, these tagged fish exhibited strong fidelity and specificity to PIE. For example, across multiple years, tagged striped bass resided in PIE waters for an average of 1.5–2.5 months per year (means: 51–72 days; range 2–122 days), left this estuary in fall, then returned in subsequent years. Third, this specificity and fidelity connected PIE to other locations. The fish exported nutrients and energy to at least three other coastal locations through biomass added as growth. These results demonstrate that what happens in an individual estuary can affect other estuaries. Striped bass that use tightly connected routes to feed in specific estuaries should have greater across-system impacts than fish that are equally likely to go anywhere. Consequently, variations in when, where, and how fish migrate can alter across-estuary impacts.

Description: We thank the Plum Island Ecosystems LTER program (OCE-0423565, OCE- 1058747, OCE-1238212) and the University of Massachusetts Intercampus Marine Sciences Graduate Program for support.