Fleeger
John W.
Fleeger
John W.
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PreprintThe response of nematodes to deep-sea CO2 sequestration : a quantile regression approach( 2010-01) Fleeger, John W. ; Johnson, David S. ; Carman, K. R. ; Weisenhorn, Pamela B. ; Gabriele, A. ; Thistle, D. ; Barry, James P.One proposed approach to ameliorate the effects of global warming is sequestration of the greenhouse gas CO2 in the deep sea. To evaluate the environmental impact of this approach, we exposed the sediment-dwelling fauna at the mouth of the Monterey Submarine Canyon (3262 m) and a site on the nearby continental rise (3607 m) to CO2- rich water. We measured meiobenthic nematode population and community metrics after ~30-day exposures along a distance gradient from the CO2 source and with sediment depth to infer the patterns of mortality. We also compared the nematode response with that of harpacticoid copepods. Nematode abundance, average sediment depth, tail-group composition, and length: width ratio did not vary with distance from the CO2 source. However, quantile regression showed that nematode length and diameter increased in close proximity to the CO2 source in both experiments. Further, the effects of CO2 exposure and sediment depth (nematodes became more slender at one site, but larger at the other, with increasing depth in the sediment) varied with body size. For example, the response of the longest nematodes differed from those of average length. We propose that nematode body length and diameter increases were induced by lethal exposure to CO2-rich water and that nematodes experienced a high rate of mortality in both experiments. In contrast, copepods experienced high mortality rates in only one experiment suggesting that CO2 sequestration effects are taxon specific.
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PreprintSusceptibility of salt marshes to nutrient enrichment and predator removal( 2006-03-15) Deegan, Linda A. ; Bowen, Jennifer L. ; Drake, Deanne C. ; Fleeger, John W. ; Friedrichs, Carl T. ; Galvan, Kari A. ; Hobbie, John E. ; Hopkinson, Charles S. ; Johnson, J. Michael ; Johnson, David S. ; LeMay, Lynsey E. ; Miller, Erin ; Peterson, Bruce J. ; Picard, Christian ; Sheldon, Sallie ; Sutherland, Michael ; Vallino, Joseph J. ; Warren, R. ScottThe 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.
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PreprintTop-down and bottom-up control of infauna varies across the saltmarsh landscape( 2007-12) Fleeger, John W. ; Johnson, David S. ; Galvan, Kari A. ; Deegan, Linda A.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.
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PreprintResponse of the benthic food web to short- and long-term nutrient 1 enrichment in saltmarsh mudflats( 2013-01) Pascal, Pierre-Yves ; Fleeger, John W. ; Boschker, Henricus T. S. ; Mitwally, Hanan M. ; Johnson, David S.We examined the responses of biota at or near the base of the benthic food web to nutrient enrichment in salt marsh mudflats in Plum Island estuary (Massachusetts, USA). To simulate eutrophication, nitrate and phosphate loading rates were increased 10- to 15-fold in creeks fertilized for 2 mo (i.e. short-term enrichment) or 6 yr (chronic enrichment). We found that benthic invertebrate community structure was not altered by nutrient enrichment, although the abundance of epifaunal, but not infaunal, grazers increased. Short-term enrichment had no effect on the food web, but significant changes were detected with chronic enrichment. Grazing experiments with 15N-enriched bacteria and 13C-enriched benthic algae revealed higher per capita ingestion rates of benthic microalgae by nematodes, copepods and hydrobiid snails in the creek with chronic nutrient enrichment where isotope composition also indicated that algae increased in dietary importance. The fraction of bacterial biomass grazed was not affected by nutrient enrichment; however, the fraction of benthic algal biomass grazed increased by 235% with chronic enrichment. This higher grazing pressure was partly the result of dietary changes (increases in per capita feeding rate or a change in selection) but was mostly due to an increased abundance of the grazing consumer with the highest biomass, the snail Nassarius obsoletus. This increased top-down control partially masked the bottom-up effects of nutrient enrichment on algal biomass and helps explain the slow and inconsistent response of microalgal biomass to chronic nutrient enrichment previously observed in this estuary. Our research shows that eutrophication may subtly affect benthic food webs before large, sustained increases in algal biomass are observed.
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PreprintMummichog Fundulus heteroclitus responses to long-term, whole-ecosystem nutrient enrichment( 2013-05) Lockfield, Konner C. ; Fleeger, John W. ; Deegan, Linda A.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.