Publications

Salt marshes are valuable ecosystems, and there is concern that increases in the rate of sea level rise along with anthropogenic activities are leading to the loss of vegetated habitat. The area of vegetated marsh can change not only through advance and retreat of the open fetch edge, but also due to channel widening and contracting, formation and drainage of interior ponds, formation and revegetation of interior mud flats, and marsh migration onto upland areas, each of which is influenced by different processes. This study used historical aerial photographs to measure changes in the extent of vegetated marsh over approximately 70 years at study marshes located in three long-term ecological research (LTER) sites along the US East coast: Georgia Coastal Ecosystems (GCE), Virginia Coast Reserve (VCR), and Plum Island Ecosystems (PIE). Marsh features were categorized into vegetated marsh, ponds, interior mud flats, and channels for three time periods at each site. The three sites showed different patterns of change in vegetated marsh extent over time. At the GCE study site, losses in vegetated marsh, which were primarily due to channel widening, were largely offset by channel contraction in other areas, such that there was little to no net change over the study period. The study marsh at VCR experienced extensive vegetated marsh loss to interior mud flat expansion, which occurred largely in low-lying areas. However, this loss was counterbalanced by marsh gain due to migration onto the upland, resulting in a net increase in vegetated marsh area over time. Vegetated marsh at PIE decreased over time due to losses from ponding, channel widening, and erosion at the open fetch marsh edge. Digital elevation models revealed that the vegetated areas of the three marshes were positioned at differing elevations relative to the tidal frame, with PIE at the highest and VCR at the lowest elevation. Understanding the patterns of vegetation loss and gain at a given site provides insight into what factors are important in controlling marsh dynamics and serves as a guide to potential management actions for marsh protection.

Keystone species have large ecological effects relative to their abundance and have been identified in many ecosystems. However, global change is pervasively altering environmental conditions, potentially elevating new species to keystone roles. Here, we reveal that a historically innocuous grazer—the marsh crab Sesarma reticulatum—is rapidly reshaping the geomorphic evolution and ecological organization of southeastern US salt marshes now burdened by rising sea levels. Our analyses indicate that sea-level rise in recent decades has widely outpaced marsh vertical accretion, increasing tidal submergence of marsh surfaces, particularly where creeks exhibit morphologies that are unable to efficiently drain adjacent marsh platforms. In these increasingly submerged areas, cordgrass decreases belowground root:rhizome ratios, causing substrate hardness to decrease to within the optimal range for Sesarma burrowing. Together, these bio-physical changes provoke Sesarma to aggregate in high-density grazing and burrowing fronts at the heads of tidal creeks (hereafter, creekheads). Aerial-image analyses reveal that resulting “Sesarma-grazed” creekheads increased in prevalence from 10 ± 2% to 29 ± 5% over the past <25 y and, by tripling creek-incision rates relative to nongrazed creekheads, have increased marsh-landscape drainage density by 8 to 35% across the region. Field experiments further demonstrate that Sesarma-grazed creekheads, through their removal of vegetation that otherwise obstructs predator access, enhance the vulnerability of macrobenthic invertebrates to predation and strongly reduce secondary production across adjacent marsh platforms. Thus, sea-level rise is creating conditions within which Sesarma functions as a keystone species that is driving dynamic, landscape-scale changes in salt-marsh geomorphic evolution, spatial organization, and species interactions.

In the nutrient-rich region surrounding marine phytoplankton cells, heterotrophic bacterioplankton transform a major fraction of recently fixed carbon through the uptake and catabolism of phytoplankton metabolites. We sought to understand the rules by which marine bacterial communities assemble in these nutrient-enhanced phycospheres, specifically addressing the role of host resources in driving community coalescence. Synthetic systems with varying combinations of known exometabolites of marine phytoplankton were inoculated with seawater bacterial assemblages, and communities were transferred daily to mimic the average duration of natural phycospheres. We found that bacterial community assembly was predictable from linear combinations of the taxa maintained on each individual metabolite in the mixture, weighted for the growth each supported. Deviations from this simple additive resource model were observed but also attributed to resource-based factors via enhanced bacterial growth when host metabolites were available concurrently. The ability of photosynthetic hosts to shape bacterial associates through excreted metabolites represents a mechanism by which microbiomes with beneficial effects on host growth could be recruited. In the surface ocean, resource-based assembly of host-associated communities may underpin the evolution and maintenance of microbial interactions and determine the fate of a substantial portion of Earth’s primary production.

Tidal freshwater marshes can protect downstream ecosystems from eutrophication by intercepting excess nutrient loads, but recent studies in salt marshes suggest nutrient loading compromises their structural and functional integrity. Here, we present data on changes in plant biomass, microbial biomass and activity, and soil chemistry from plots in a tidal freshwater marsh on the Altamaha River (GA) fertilized for 10 yr with nitrogen (+N), phosphorus (+P), or nitrogen and phosphorus (+NP). Nitrogen alone doubled aboveground biomass and enhanced microbial activity, specifically rates of potential nitrification, denitrification, and methane production measured in laboratory incubations. Phosphorus alone increased soil P and doubled microbial biomass but did not affect microbial processes. Nitrogen or P alone decreased belowground biomass and soil carbon (C) whereas +NP increased aboveground biomass, microbial biomass and N cycling, and N, P, and C assimilation and burial more than either nutrient alone. Our findings suggest differential nutrient limitation of tidal freshwater macrophytes by N and microbes by P, similar to what has been observed in salt marshes. Macrophytes outcompete microbes for P in response to long‐term N and P additions, leading to increased soil C storage through increased inputs of belowground biomass relative to N and P added singly. The susceptibility of tidal freshwater marshes to long‐term nutrient enrichment and, hence their ability to mitigate eutrophication will depend on the quantity and relative proportion of N vs. P entering estuaries and tidal wetlands.

Benthic foraminifera are valuable indicators in environmental studies, including those on marine pollution monitoring. While a great deal of foraminiferal biomonitoring research utilizes abundance and distributional data, further value resides in better understanding the incorporation of heavy metal pollutants in foraminiferal calcite. By experimentally growing assemblages of foraminifera from propagules (small juveniles) gathered from Sapelo Island, Georgia and Little Duck Key, Florida, this study examines foraminiferal incorporation of the heavy metals arsenic, cadmium, nickel, and zinc over a range of concentrations.

Surface sediment was collected and sieved to concentrate the propagules. The propagules were then used to experimentally grow assemblages with each exposed to a different heavy metal. After one month, the experimentally grown foraminifera were harvested and samples of the two most common species from each location, Ammonia tepida (Cushman) and Haynesina germanica (Ehrenberg) from Sapelo Island and Quinqueloculina sabulosa (Cushman) and Triloculina oblonga (Montagu) from Little Duck Key, were selected for trace element analysis. Calcite of the tests was analyzed using LA-ICP-MS to quantify the heavy metal incorporation.

Rotalid species A. tepida and H. germanica incorporated more cadmium as its concentration in the surrounding water increased, whereas miliolid species Q. sabulosa and T. oblonga incorporated more of the metals zinc and nickel. This study shows that while foraminiferal incorporation of heavy metals has great potential as a biomonitoring tool, multiple factors (especially inter-clade variation) must be considered carefully. In future marine environmental research, these factors may help to create a more targeted assessment of environmental pollution.

The eastern oyster (Crassostrea virginica) is an important proxy for examining historical trajectories of coastal ecosystems. Measurement of ~40,000 oyster shells from archaeological sites along the Atlantic Coast of the United States provides a long-term record of oyster abundance and size. The data demonstrate increases in oyster size across time and a nonrandom pattern in their distributions across sites. We attribute this variation to processes related to Native American fishing rights and environmental variability. Mean oyster length is correlated with total oyster bed length within foraging radii (5 and 10 km) as mapped in 1889 and 1890. These data demonstrate the stability of oyster reefs despite different population densities and environmental shifts and have implications for oyster reef restoration in an age of global climate change.

Aim: Introduced species may display or foster novel latitudinal clines because they are not well adapted to their new habitats. We tested the hypothesis that the latitudinal cline in nematode diversity in salt marshes would differ between the native (United States) and introduced (China) ranges of Spartina alterniflora.
Location: East Coasts of the United States (30.32–43.33°N) and China
(30.36–39.14°N).
Methods: We extracted nematodes from soil samples collected at 32 sites along the United States East Coast and 41 sites along the Chinese coast. We compared latitudinal patterns in nematode diversity and composition between the native and introduced ranges.
Results: In the native range of S. alterniflora, nematode richness at lower latitudes was almost twice as high as that at higher latitudes. In contrast, we found no latitudinal pattern in nematode richness or diversity in the introduced range of S. alterniflora. Nematode genus richness at all sites in China was about half that at lower latitudes in the United States. Beta diversity of nematodes increased with geographic distance in the United States, but not China.
Main conclusions: Nematode diversity did not show latitudinal clines in salt marshes dominated by introduced S. alterniflora in China. A likely explanation is that the recently introduced populations are still relatively genetically homogenous, whereas in the native range, genetic variation in plant populations across latitude drives different nematode communities. We suggest that future studies of introduced species will gain additional insights by taking an explicitly geographic perspective.

Elevation differences in salt marshes result in numerous ecological consequences as a result of variation in tidal flooding. We demonstrate here that elevation differences are also negatively correlated with soil temperature on the marsh platform, irrespective of tidal flooding. Field observations of soil temperature at 10‐cm depth in a Georgia marsh showed that elevation increases of 0.5 m corresponded to decreases in average soil temperature of 0.9–1.7°C during both winter and summer. Landsat 8 estimates of land surface temperatures across the marsh in dry (nonflooded) scenes also showed that temperature decreased with increasing elevation, which was consistent with soil observations. Similar satellite results were also found in a test marsh in Virginia. Biological reactions are temperature‐dependent, and these findings indicate that metabolic processes will vary over short distances. This is important for accurately estimating marsh metabolism and predicting how changes in temperature will affect future productivity.

Climate change is altering consumer−plant interactions in ecosystems worldwide. How consumers alter their spatial distribution, grazing activities, and functional morphology in response to climate stress can determine whether their effects on plants intensify or relax. Few studies have considered multiple
consumer response metrics to elucidate the mechanisms underpinning the resulting changes in consumer−plant interactions. Here, we tested how drought stress influences the interaction between the dominant consumer, the fungal-farming periwinkle snail Littoraria irrorata, and a foundational plant, cordgrass Spartina alterniflora, in a southeastern US salt marsh. In a 4 mo
field experiment, we maintained moderate snail densities in mesh control chambers and clear plastic climate chambers that simulated drought by elevating temperatures and drying soils. Monitoring revealed that snails
more often congregated on cordgrass stems than leaves in climate chambers than in controls. Image analyses indicated that this behavioral shift corresponded to snails inflicting shorter, but more numerous, fungal-infested scars on cordgrass leaves, and causing less plant damage in climate chambers than controls. Coincident with their net reduction in grazing, snails maintained longer radulae, whose central teeth were blunter and lateral teeth were sharper, in climate chambers compared to controls. These results suggest that under drought, snail radulae may experience less frictional wear and that, at inter mediate densities, snail−cordgrass interactions re lax. Together with prior
research showing that at high densities, snails can denude cordgrass during drought, we conclude that consumer density, behavior, and morphological responses must be integrated in predictions of how climate change will affect the direction, strength, and stability of consumer−plant interactions.

1. Salt marshes suffered large‐scale degradation in recent decades. Extreme events such as hot and dry spells contributed significantly to this, and are predicted to increase not only in intensity, but also in frequency under future climate scenarios. Such repetitive extreme events may generate cumulative effects on ecosystem resilience. It is therefore important to elucidate how marsh vegetation responds to repetitive stress, and whether changes in key species interactions can modulate vegetation resilience.
2. In this study, we investigated how moderate but repetitive desiccation events, caused by the combined effects of drought and high temperatures, affect cordgrass (Spartina alterniflora), the dominant habitat‐forming grass in southeastern US salt marshes. In a 4‐month field experiment, we simulated four consecutive desiccation events by periodically excluding tidal flooding and rainfall, while raising temperature. We crossed this desiccation treatment with the presence/absence of ribbed mussels (Geukensia demissa) – a mutualist of cordgrass known to enhance its desiccation resilience – and with grazing pressure by the marsh periwinkle (Littoraria irrorata) that is known to suppress cordgrass’ desiccation resilience.
3. We found that each subsequent desiccation event deteriorated sediment porewater conditions, resulting in high salinity (53 ppt), low pH‐levels (3.7) and increased porewater Al and Fe concentrations (≈800 μmol/L and ≈1,500 μmol/L) upon rewetting. No effects on porewater chemistry were found as a result of snail grazing, while ribbed mussels strongly mitigated desiccation effects almost to control levels and increased cordgrass biomass by approximately 128%. Importantly, although cordgrass generally appeared healthy above‐ground at the end of the experiment, we found clear negative responses of the repetitive desiccation treatment on cordgrass below‐ground biomass, on proline (osmolyte) levels in shoots and on the number of tillers (−40%), regardless of mussel and/or snail presence.
4. Synthesis. Even though the mutualism with mussels strongly mitigated chemical effects in the sediment porewater throughout the experiment, mussels could not buffer the adverse ecophysiological effects observed in cordgrass tissue. Our results therefore suggest that although mussels may alleviate desiccation stress, the predicted increased frequency and intensity of hot dry spells may eventually affect saltmarsh resilience by stressing the mutualism beyond its buffering capacity.

Restoration efforts have been escalating worldwide in response to widespread habitat degradation. However, coastal restoration attempts notoriously vary in their ability to establish resilient, high-functioning ecosystems. Conventional restoration attempts disperse transplants in competition-minimizing arrays, yet recent studies suggest that clumping transplants to maximize facilitative interactions may improve restoration success. Here, we modify the stress gradient hypothesis to generate predictions about where each restoration design will perform best across environmental stress gradients. We then test this conceptual model with field experiments manipulating transplant density and configuration across dune elevations and latitudes. In hurricane-damaged Georgia (USA) dunes, grass transplanted in competition-minimizing (low-density, dispersed) arrays exhibited the highest growth, resilience to disturbance and dune formation in low-stress conditions. In contrast, transplants survived best in facilitation-maximizing (high-density, clumped) arrays in high-stress conditions, but these benefits did not translate to higher transplant growth or resilience. In a parallel experiment in Massachusetts where dune grasses experience frequent saltwater inundation, fewer transplants survived, suggesting that there are thresholds above which intraspecific facilitation cannot overcome local stressors. These results suggest that ecological theory can be used to guide restoration strategies based on local stress regimes, maximizing potential restoration success and return-on-investment of future efforts.

Dissolved organic matter (DOM) is a large and complex mixture of compounds with source inputs that differ with location, season and environmental conditions. Here, we investigated drivers of DOM composition changes in a marsh‐dominated estuary off the southeastern U.S. Monthly water samples were collected at a riverine and estuarine site from September 2015 to September 2016, and bulk, optical, and molecular analyses were conducted on samples before and after dark incubations. Results showed that river discharge was the primary driver changing the DOM composition at the mouth of the Altamaha River. For discharge higher than ~ 150 m³ s^‐1, DOC concentrations and the terrigenous character of the DOM increased approximately linearly with river flow. For low discharge conditions, a clear signature of salt marsh‐derived compounds was observed in the river. At the head of Sapelo Sound, changes in DOM composition were primarily driven by river discharge and possibly by summer algae blooms. Microbial consumption of DOC was larger during periods of high discharge at both sites, potentially due to the higher mobilization and influx of fresh material to the system. The Georgia coast was hit by Hurricane Matthew in October 2016, which resulted in a large input of carbon to the estuary. The DOC concentration was ~ 2 times higher and DOM composition was more aromatic with a stronger terrigenous signature compared to the seasonal maximum observed earlier in the year during peak river discharge conditions. This suggests that extreme events notably impact DOM quantity and quality in estuarine regions.

In estuaries, future variation in sea level and river discharge will lead to saline intrusion into low-salinity tidal marshes. To investigate the processes that control the differential response and recovery of tidal freshwater marsh plant communities to saline pulses, a 3 × 5 factorial greenhouse experiment was conducted to examine the effects of a range of salinity levels (3, 5, and 10 practical salinity units (PSU)) and pulse durations (5, 10, 15, 20, and 30 days per month) on community composition of tidal freshwater marsh vegetation. Recovery of perturbed communities was also examined after 10 months. The results showed that community composition was increasingly affected by the more-saline and longer-duration treatments. The increasing suppression of salt-sensitive species resulted in species reordering, decreased species richness, and decreased aboveground biomass. Most of the plant species were able to recover from low-salinity, short-duration saline pulses in less than 1 year. However, because not all species recovered in the heavily salinized treatments, species richness at the end of the recovery period remained low for treatments that were heavily salinized during the treatment period. In contrast, plant aboveground biomass fully recovered in the heavily salinized treatments. Although the magnitude and duration of pulsed environmental changes had strong effects on community composition, shifts in community composition prevented long-term reductions in productivity. Thus, in this study system, environmental change affected species composition more strongly than it did ecosystem processes.

1. Plants adjust their size and reproductive effort in response to numerous selection pressures and constraints. The self‐thinning law describes a well‐known trade‐off between size and density. Plants also trade‐off investment into growth vs. sexual reproduction, as described by life‐history theory.
2. We build on past work on plant allometry and life history by examining both self‐thinning and size‐dependent reproduction in a single plant species, the saltmarsh grass Spartina alterniflora, across a wide range of settings: three landscape positions, two habitats and eight sites, across sixteen years.
3. Plants in different landscape positions and years varied tremendously in size and shoot density. However, all this variation could be explained by a single allometric relationship consistent with the self‐thinning law, but with a lower slope. Flowering was size‐dependent, and the size at which plants had a 50% probability of flowering varied among habitat, sites and years. Plants that were stressed reproduced at a smaller size than plants that were growing under good conditions, and this pattern was consistent among habitat, sites and years. Finally, reproductive biomass and the proportion of shoots flowering increased with increasing vegetative size (plant height or shoot biomass). Combining these two patterns, S. alterniflora plants growing high density are small and reproduce at a smaller size than large plants growing at low density.
4. Although there is tremendous spatial and temporal variation in S. alterniflora growth and reproductive patterns, all this variation can be understood as resulting from two simple allometric trade‐offs. Because saltmarsh plants often occur in monospecific stands, they may serve as simple, model systems for studies of plant life history.

Phenology studies mostly focus on variation across time or landscapes. However, phenology can vary at fine spatial scales, and these differences may be as important as long-term change from climate warming. We used high frequency “PhenoCam” data to examine phenology of Spartina alterniflora, a foundation species native to salt marshes on the US East and Gulf coasts, and a common colonizer elsewhere. We examined phenology across three microhabitats from 2013 to 2017 and used this information to create the first spring green-up model for S. alterniflora. We then compared modern spatial variation to that exhibited over a 60-year climate record. Marsh interior plants initiated spring growth 17 days earlier than channel edge plants and spent 35 days more in the green-up phenophase and 25 days less in the maturity phenophase. The start of green-up varied by 17 days among 3 years. The best spring green-up model was based on winter soil total growing degree days. Across microhabitats, spring green-up differences were caused by small elevation changes (15 cm) that drove soil temperature variation of 0.8°C. Preliminary evidence indicated that high winter belowground biomass depletion triggered early green-up. Long-term change was similar: winter soil temperatures warmed 1.7 ± 0.3°C since 1958, and green-up advanced 11 ± 6 days, whereas contemporary microhabitat differences were 17 ± 4 days. Incorporating local spatial variation into plant phenology models may provide an early warning of climate vulnerability and improve understanding of ecosystem-scale productivity. Microscale phenology variation likely exists in other systems and has been unappreciated.

Exchange of groundwater is an important transfer mechanism for nutrients and pollutants between coastal aquifers and surface waters. Constraining such exchange in salt marshes – where biological productivity and biogeochemical cycling rates are among the highest of all coastal ecosystems – is vital for understanding ecosystem function and vulnerability. Here, we quantify groundwater discharge into the tidal Duplin River from the adjoining salt marsh near Sapelo Island, Georgia using high spatial and temporal resolution field measurements of radon-222. Field campaigns occurred for several weeks each summer during 2013, 2015, and 2016. Spatial surveys reveal a general increase in radon activity upstream through the Duplin River, which may result from either higher groundwater discharge or lower mixing rates in the headwaters. To distinguish between these possibilities, we use a radon mass balance model to determine groundwater input. We find that groundwater discharge (normalized to inundated marsh surface area) to the headwaters average 5.1–5.8 cm³/cm²marsh/day across all three field campaigns, which are comparable to those to the main channel (averaging 6.0–6.5 cm³/cm² marsh/day across all three field campaigns). Our work reveals a positive relationship between aerial extent of marsh inundation and groundwater discharge into the Duplin River. Discharge is generally maximal during falling tide, reflecting a hydraulic gradient driver, but also is significant prior to high tide, indicative of sediment compression as a driver of groundwater inputs. Constraining the relationship between marsh inundation and resulting groundwater dynamics is an integral aspect to assessing how salt marsh circulation processes may respond to intensifying inundation (from reduced sediment supply, subsidence, and/or rising sea levels).

Benthic foraminifera have long served as indicators of environmental conditions – both natural and anthropogenically impacted. To better understand the responses of benthic foraminifera to specific heavy metal contaminants (Cd, Pb, Zn), assemblages of coastal benthic foraminifera were grown from propagules (tiny juveniles) in the lab with exposure to a single heavy metal over a range of concentrations, based on the US Environmental Protection Agency’s Critical Maximum Concentration (CMC) values. Foraminiferal propagule banks were collected from relatively pristine mudflats, located on the southern end of Sapelo Island, Georgia (USA). Consistent with the findings of numerous field-based studies, foraminifera were found to respond negatively to Cd, Pb, and Zn. Overall, assemblages grown with exposure to higher concentrations of these metals are characterized by decreased abundances, species richness, and evenness. All of the acute responses observed in these metrics occur at concentrations equal to or somewhat higher than the USEPA’s CMC values. Foraminiferal responses vary by metal, though the four most common species (two monothalamids: Ovammina opacaDahlgren, Psammophaga sapela Altin Ballero, Habura, Goldstein; and two rotaliids: Haynesina germanica (Ehrenberg), Ammonia tepida (Cushman)) responded in a broadly similar fashion. The monothalamid species however may be more sensitive to high concentrations of each metal. Of the metals examined, exposure to Pb had the most deleterious effect, followed by Zn, then Cd. These four most abundant species appear to be more tolerant of Cd than the other metals. Zn was the only metal in the study that produced abundant aberrant test morphologies. Ammonia tepida grew abnormally more frequently than any other species encountered and exhibited a distinctive enlarged aperture as well as aberrant patterns of calcification, chamber arrangement, and enlarged pores. Abnormal tests were also found in H. germanica and a miliolid. The monothalamid species did not produce aberrant test morphologies. Results support the application of foraminifera as bio-indicators in polluted environments.

Habitat patch composition and configuration mediate the fitness and distribution of many species. However, we know little about how this landscape complementation may influence the distribution of an invasive species’ ecological impacts and, in turn, how this affects ecosystem resilience to disturbance. We surveyed > 820 km of coastline to evaluate how landscape complementation mediates patterns in invasive feral hog (Sus scrofa) rooting, trampling and wallowing disturbances in southeastern US salt marshes and assessed marsh resilience to these behaviors in an 8-site survey and 13-month field experiment. We discovered that hog rooting and trampling most often occur where hardwood forest comprises > 30% and salt marsh < 22% of habitat surrounding each surveyed site, respectively, while wallowing correlated most strongly with salt marsh invertebrate densities. At the 8 survey sites, vegetation cover, soil organic carbon, and surface elevation were consistently lower, and soil anoxia and porewater ammonium-nitrogen higher, in hog-disturbed relative to undisturbed areas. The experiment revealed that vegetation can recover when rooted or trampled, but remains depressed when wallowed or repeatedly disturbed. Together, these findings provide novel evidence that habitat patch composition at landscape scales can act together with local habitat attributes to dictate invasive species’ disturbance patterns and highlight areas most vulnerable to invaders. In salt marshes, insights gleaned from such consideration of landscape complementation can inform conservation and management strategies for curbing the impact of this prolific, global invader.

Benthic foraminifera are valuable environmental indicators of heavy metal contaminants in marine environments. To broaden their effectiveness as bioindicators, this study compares individually the effects of selected heavy metal contaminants, including both metabolically essential and non-essential elements, on temperate rotalids and subtropical miliolids, as well as associated monothalamid foraminifera. To accomplish these aims, assemblages of foraminifera were grown experimentally from propagules (small juveniles) collected from two coastal sites: Sapelo Island, Georgia, and Little Duck Key, Florida, that provide an effective comparison between environments and types of foraminifera. Surface sediment was collected from both locations and sieved immediately after collection. Using the propagule method, assemblages of foraminifera were grown in the laboratory from propagules in the sediment samples. Two metabolically essential trace elements, nickel, and zinc, and two non-essential elements, arsenic and cadmium were used to represent both types of heavy metal. Experimental conditions were held constant while varying only the metal concentrations.

In treatments from both origins, increasing concentrations of cadmium, nickel, and zinc led to decreases in abundance and diversity for the foraminifera. In addition, zinc, and to a lesser extent cadmium and nickel above certain concentrations, resulted in an increase of deformed tests among the foraminifera. Deformities occurred amongst the most common calcareous species from Sapelo island: Ammonia tepida and Haynesina germanica. Fewer deformities were observed in common calcareous species from Little Duck Key, the miliolids Quinqueloculina sabulosa and Quinqueloculina bosciana featured few deformities. Notably, monothalamid species such as Psammophaga sapelaremained present at high metal concentrations. These results support previous research and reinforce the usefulness of rotalids such as A. tepida and H. germanica as bioindicators of heavy metal contamination as well as suggesting a possible use of monothalamids such as P. sapela in this manner.

Sea level rise is expected to increase inundation and saltwater intrusion into many tidal freshwater marshes and forests. Saltwater intrusion may be long-term, as with rising seas, or episodic, as with low river flow or storm surge. We applied continuous (press) and episodic (pulse) treatments of dilute seawater to replicate 2.5 × 2.5 m field plots for three years and measured soil attributes, including soil porewater, oxidation-reduction potential, soil carbon (C), and nitrogen (N) to investigate the effects of continuous and episodic saltwater intrusion and increased inundation on tidal freshwater marsh elemental cycling and soil processes. Continuous additions of dilute seawater resulted in increased porewater chloride, sulfate, sulfide, ammonium, and nitrate concentrations. Plots that received press additions also had lower soil oxidation-reduction potentials beginning in the second year. Episodic additions of dilute seawater during typical low flow conditions (Sept.-Oct.) resulted in transient increases in porewater chloride and sulfate that returned to baseline conditions once dosing ceased. Freshwater additions did not affect porewater inorganic N or soil C or N. Persistent saltwater intrusion in freshwater marshes alters the N cycle by releasing ammonium-N from sorption sites, increasing nitrification and severely reducing N storage in macrophyte biomass. Chronic saltwater intrusion, as is expected with rising seas, is likely to shift tidal freshwater marshes from a sink to a source of N whereas intermittent intrusion from drought may have no long term effect on N cycling.

Tidal salt marshes sequester and store blue carbon at both short and long time scales. Marsh soils shape and maintain the ecosystem by supporting complex biogeochemical reactions, deposition of sediment, and accumulation of organic matter. In this study, we examined the potential of imaging spectroscopy techniques to indirectly quantify and map tidal marsh soil properties at a National Estuarine Research Reserve in Georgia, USA. A framework was developed to combine modern digital image processing techniques for marsh soil mapping, including object-based image analysis (OBIA), machine learning modeling, and ensemble analysis. We also evaluated the efficacy of airborne hyperspectral sensors in estimating marsh soil properties compared to spaceborne multispectral sensors, WorldView-2 and QuickBird. The pros and cons of object-based modeling and mapping were assessed and compared with traditional pixel-based mapping methods. The results showed that the designed framework was effective in quantifying and mapping three marsh soil properties using the composite reflectance from salt marsh environment: soil salinity, soil water content, and soil organic matter content. Multispectral sensors were successful in quantifying soil salinity and soil water content but failed to model soil organic matter. The study also demonstrated the value of minimum noise fraction transformation and ensemble analysis techniques for marsh soil mapping. The results suggest that imaging spectroscopy based modeling is a promising tool to quantify and map marsh soil properties at a local scale, and is a potential alternative to traditional soil data acquisition to support carbon cycle research and the conservation and restoration of tidal marshes.

Climate change and consumer outbreaks are driving ecosystem collapse worldwide. Although much research has demonstrated that these factors can interact, how heterogeneity in top–down control intensity and physical forcing modulates ecosystem resilience to climate stress remains poorly understood. Here, we explore whether the nocturnal herbivorous crab Sesarma reticulatum can control spatially dominant cordgrass (Spartina alterniflora) growth and how its top–down effects vary with crab density, drought stress, and large-scale disturbance in southeastern US salt marshes. In multiple field experiments and surveys, we show that Sesarma depresses cordgrass growth and that its effects increase in a saturating manner with increasing crab density, such that the highest naturally occurring densities of this consumer can trigger local cordgrass die-off. This top–down effect of Sesarma is similar in magnitude to what is thought to be the dominant grazer in the system, the marsh periwinkle snail Littoraria irrorata. In a drought stress by Sesarma density experiment, we further show that salinity stress and intensive crab herbivory additively suppress cordgrass drought resistance. After drought subsides, surveys and experiments reveal that Sesarma also stifles cordgrass re-growth into existing die-off areas. Together, these results show that multiple grazers powerfully regulate the productivity and drought resilience of these intertidal grasslands and that heterogeneity in physical stress and consumer density can dictate when and where top–down forcing is important. More generally, this work provides a rare, experimental demonstration of the critical role top–down control can play across the initiation and recovery stages of ecosystem die-off.

Knowledge of light partitioning into different optically active constituents, particularly chromophoric dissolved organic matter (CDOM) in the ultraviolet (UV) is indispensable for understanding UV dependent biogeochemical issues including photochemical processes in optically complex waters. Herein a new approach is presented to investigate photochemistry by blending two ocean color algorithms, namely the composite SeaUV (Cao et al., 2014) and the SeaCDOM (Cao and Miller, 2015) algorithms, and applying them to visible remote sensing reflectance (Rrs) measured using the Hyperspectral Imager for the Coastal Ocean (HICO). As illustrated using photochemical carbon monoxide (CO) production from CDOM, this model approach allows high resolution examination of UV optical details with estimates of both depth-specific and depth-integrated photoproduction rates in a dynamic estuarine/coastal environment. Decoupled retrievals of inherent and apparent optical properties (i.e. diffuse attenuation coefficient (K_d) and CDOM absorption coefficient (a_g)) using two distinct ocean color algorithms over the entire UV spectrum allow a synoptically dynamic view of CDOM’s contribution to light attenuation (a_g/K_d). This provides new potential to probe UV processes in complex coastal waters on regional as well as global scales using remote sensing of ocean color.

Dissolved organic nitrogen (DON) can account for a large fraction of the dissolved nitrogen (N) pool in the ocean, but the cycling of marine DON is poorly understood. Recent discoveries that urea‐ and cyanate‐N can be oxidized by some strains of Thaumarchaeota suggest that these abundant microbes may be able to access and oxidize a fraction of the DON pool. However, measurements of the oxidation of N supplied as DON compounds are scarce. Here, we compare oxidation rates of N supplied as a variety of DON compounds in samples from Georgia coastal waters, where nitrifier communities are numerically dominated by Thaumarchaeota. Our data indicate that polyamine‐N is particularly amenable to oxidation compared to the other DON compounds tested. Oxidation of N supplied as putrescine (1,4‐diaminobutane) was generally higher than that of N supplied as glutamate, arginine, or urea, and was consistently 5–10% of the ammonia oxidation rate. Our data also suggest that the oxidation rate of polyamine‐N may increase as the length of the carbon skeleton increases. Oxidation of N supplied as putrescine, urea, and glutamate were all highest near the coast and lower further offshore, consistent with patterns of ammonia oxidation in these waters. Though it is unclear whether oxidation of polyamine‐N reflects direct oxidation by Thaumarchaeota or combines remineralization and subsequent ammonia oxidation, more rapid oxidation of N from putrescine compared to amino acids or urea suggests that polyamine‐N may contribute significantly to nitrification in the ocean.

Small-scale armoring placed near the marsh-upland interface to protect single-family homes is widespread but understudied. Using a nested, spatially blocked sampling design on the coast of Georgia, USA, we compared the biota and environmental characteristics of 60 marshes adjacent to either a bulkhead, a residential backyard with no armoring, or an intact forest. We found that marshes adjacent to bulkheads were at lower tidal elevations and had features typical of lower elevation marsh habitats: high coverage of the marsh grass Spartina alterniflora, high density of crab burrows, and muddy sediments. Marshes adjacent to unarmored residential sites had higher soil water content and lower porewater salinities than the armored or forested sites, suggesting that there may be increased freshwater input to the marsh at these sites. Deposition of Spartina wrack on the marsh-upland ecotone was negatively related to elevation at armored sites and positively related at unarmored residential and forested sites. Armored and unarmored residential sites had reduced densities of the high marsh crab Armases cinereum, a species that moves readily across the ecotone at forested sites, using both upland and high marsh habitats. Distance from the upland to the nearest creek was longest at forested sites. The effects observed here were subtle, perhaps because of the small-scale, scattered nature of development. Continued installation of bulkheads in the southeast could lead to greater impacts such as those reported in more densely armored areas like the northeastern USA. Moreover, bulkheads provide a barrier to inland marsh migration in the face of sea level rise. Retaining some forest vegetation at the marsh-upland interface and discouraging armoring except in cases of demonstrated need could minimize these impacts.

Cascading consequences of predator extinctions are well documented, but impacts of perturbations to predator size‐structure and how these vary across species remain unclear. Body size is hypothesized to be a key trait governing individual predators’ impact on ecosystems. Therefore, shifts in predator size-structure should trigger ecosystem ramifications which are consistent across functionally similar species. Using a US salt marsh as a model system, we tested this hypothesis by manipulating size class (small, medium, and large) and size diversity (combination of all three size classes) within two closely related and functionally similar predatory crab species over 4 months. Across treatments, predators suppressed densities of a dominant grazer and an ecosystem engineer, enhanced plant biomass, and altered sediment properties (redox potential and saturation). Over the metabolically equivalent experimental predator treatments, small size class predators had stronger average impacts on response variables, and size class interacted with predator species identity to drive engineer suppression. Within both predator species, size diversity increased
cannibalism and slightly weakened the average impact. These results show that predator impacts in a salt marsh ecosystem are determined by both size class and size diversity; they also highlight that size class can have species‐dependent and response‐dependent effects, underlining the challenge of generalizing trait effects.

Tidal freshwater ecosystems experience acute seawater intrusion associated with periodic droughts, but are expected to become chronically salinized as sea level rises. Here we report the results from an experimental manipulation in a tidal freshwater Zizaniopsis miliacea marsh on the Altamaha River, GA where diluted seawater was added to replicate marsh plots on either a press (constant) or pulse (2 months per year) basis. We measured changes in porewater chemistry (SO₄²⁻, Cl⁻, organic C, inorganic nitrogen and phosphorus), ecosystem CO₂ and CH₄exchange, and microbial extracellular enzyme activity. We found that press (chronic) seawater additions increased porewater chloride and sulfate almost immediately, and ammonium and phosphate after 2–4 months. Chronic increases in salinity also decreased net ecosystem exchange, resulting in reduced CO₂ and CH₄ emissions from press plots. Our pulse treatment, designed to mimic natural salinity incursion in the Altamaha River (September and October), temporarily increased porewater ammonium concentrations but had few lasting effects on porewater chemistry or ecosystem carbon balance. Our findings suggest that long-term, chronic saltwater intrusion will lead to reduced C fixation and the potential for increased nutrient (N, P) export while acute pulses of saltwater will have temporary effects.

Coastal low-salinity marshes are increasingly experiencing periodic to extended periods of elevated salinities due to the combined effects of sea level rise and altered hydrological and climatic conditions. However, we lack the ability to predict detailed vegetation responses, especially for saline pulses that are more realistic in nature than permanent saline presses. In this study, we exposed common freshwater and brackish plants to different durations (1–31 days per month for 3 months) of saline water (salinity of 5). We found that Zizaniopsis miliacea was more tolerant to salinity than the other two freshwater species, Polygonum hydropiperoides and Pontederia cordata. We also found that Zizaniopsis miliaceabelowground and total biomass appeared to increase with salinity pulses up to 16 days in length, although this relationship was quite variable. Brackish plants, Spartina cynosuroides, Schoenoplectus americanus and Juncus roemerianus, were unaffected by the experimental treatments. Our experiment did not evaluate how competitive interactions would further affect responses to salinity but our results suggest the hypothesis that short pulses of saline water will increase the cover of Zizaniopsis miliacea and decrease the cover of Polygonum hydropiperoides and Pontederia cordata in tidal freshwater marshes, thereby reducing diversity without necessarily affecting total plant biomass.

Tidal fresh marshes are at least as productive as nearby salt marshes, but much less is known about controls on primary production in tidal fresh vs. salt marshes. We studied a tidal fresh marsh in Georgia, U.S.A., dominated by the C3 grass Zizaniopsis miliacea. We documented seasonal variation in Z. miliacea above‐ground biomass and below‐ground macro‐organic matter over 1 yr, and annual variation in end‐of‐season aboveground biomass over 15 yr in creekbank and midmarsh zones. Aboveground biomass showed a distinct peak in July and October. Belowground macro‐organic matter was much greater than aboveground biomass and peaked in October. Overall productivity was similar to that of salt marshes downstream. Z. miliacea end‐of‐season above‐ground biomass showed a classic hump‐shaped “subsidy‐stress” relationship with plot elevation, but on average the creekbank supported about twofold more above‐ground biomass than the midmarsh, and both zones varied in biomass about 1.7‐fold among years. Annual variation in above‐ground biomass was negatively correlated with maximum and mean temperature in both zones, and positively with river discharge in the creekbank zone. Sea level, precipitation and water column salinity showed biologically plausible trends with respect to biomass. The responses of Z. miliacea to abiotic drivers were muted compared with the responses of nearby salt marshes dominated by Spartina alterniflora. Temperature was more important for Z. miliacea, whereas drivers of porewater salinity were more important in the salt marsh. Likely future changes in temperature, precipitation, and river discharge may pose a threat to the high productivity of tidal fresh marshes.

The outcome of species interactions may manifest differently at different spatial scales; therefore, our interpretation of observed interactions will depend on the scale at which observations are made. For example, in ladybeetle–aphid systems, the results from small‐scale cage experiments usually cannot be extrapolated to landscape‐scale field observations. To understand how ladybeetle–aphid interactions change across spatial scales, we evaluated predator–prey interactions in an experimental system. The experimental habitat consisted of 81 potted plants and was manipulated to facilitate analysis across four spatial scales. We also simulated a spatially explicit metacommunity model parallel to the experiment. In the experiment, we found that the negative effect of ladybeetles on aphids decreased with increasing spatial scales. This pattern can be explained by ladybeetles strongly suppressing aphids at small scales, but not colonizing distant patches fast enough to suppress aphids at larger scales. In the experiment, the positive effects of aphids on ladybeetles were strongest at three‐plant scale. In a model scenario where predators did not have demographic dynamics, we found, consistent with the experiment, that both the effects of ladybeetles on aphids and the effects of aphids on ladybeetles decreased with increasing spatial scales. These patterns suggest that dispersal was the primary cause of ladybeetle population dynamics in our experiment: aphids increased ladybeetle numbers at smaller scales because ladybeetles stayed in a patch longer and performed area‐restricted searches after encountering aphids; these behaviors did not affect ladybeetle numbers at larger spatial scales. The parallel experimental and model results illustrate how predator–prey interactions can change across spatial scales, suggesting that our interpretation of observed predator–prey dynamics would differ if observations were made at different scales. This study demonstrates how studying ecological interactions at a range of scales can help link the results of small‐scale ecological experiments to landscape‐scale ecological problems.

Mid-summer peaks in the abundance of Thaumarchaeota and nitrite concentration observed on the Georgia, USA, coast could result from in situ activity or advection of populations from another source. We collected data on the distribution of Thaumarchaeota, ammonia-oxidizing betaproteobacteria (AOB), Nitrospina, environmental variables and rates of ammonia oxidation during six cruises in the South Atlantic Bight (SAB) from April to November 2014. These data were used to examine seasonality of nitrification in offshore waters and to test the hypothesis that the bloom was localized to inshore waters. The abundance of Thaumarchaeota marker genes (16S rRNA and amoA) increased at inshore and nearshore stations starting in July and peaked in August at >10⁷ copies L⁻¹. The bloom did not extend onto the mid-shelf, where Thaumarchaeota genes ranged from 10³ to 10⁵ copies L⁻¹. Ammonia oxidation rates (AO) were highest at inshore stations during summer (to 840 nmol L⁻¹ d⁻¹) and were always at the limit of detection at mid-shelf stations. Nitrite concentrations were correlated with AO (R = 0.94) and were never elevated at mid-shelf stations. Gene sequences from samples collected at mid-shelf stations generated using Archaea 16S rRNA primers were dominated by Euryarchaeota; sequences from inshore and nearshore stations were dominated by Thaumarchaeota. Thaumarchaeota were also abundant at depth at the shelf-break; however, this population was phylogenetically distinct from the inshore/nearshore population. Our analysis shows that the bloom is confined to inshore waters during summer and suggests that Thaumarchaeota distributions in the SAB are controlled primarily by photoinhibition and secondarily by water temperature.

In coastal marsh ecosystems, porewater salinity strongly affects vegetation distribution and productivity. To simulate marsh porewater salinity, an integrated, spatially explicit model was developed, accounting for tidal inundation, evaporation, and precipitation, as well as lateral and vertical exchanges in both surface waters and the subsurface. It was applied to the Duplin River marsh, Sapelo Island, USA, over a 3-year period, which covered both drought and wet conditions. Simulated porewater salinity in the low and high marsh correlated with Duplin River salinity, with evapotranspiration and precipitation leading to substantial variations in porewater salinities across seasons, in particular in the high marsh. The model revealed substantial interannual variability in marsh soil conditions, and—due to its process-based approach linked to external forcings—can be used to explore effects of sea level rise and changes in hydrological forcings on marsh soil conditions.

We measured plant community composition and productivity, soil accretion, and C, N, and P burial in a tidal freshwater forest of the Altamaha River, Georgia to gain a better understanding of the ecosystem services they deliver and their ability to keep pace with current and future rates of sea level rise. Ten species were identified in two 0.1 ha plots. Nyssa aquatica (Tupelo Gum) made up 50% of the density and 57% of the total basal area. Nyssa biflora, Liquidambar styraciflua, and Fraxinus pennsylvanica were the next dominant species, collectively accounting for 37% of the density and 26% of the total basal area. Taxodium distichum only accounted for 3% of the density, but 12% of the total basal area. Aboveground productivity, measured as litterfall and stem wood growth, averaged 927 and 1030 g/m² in 2015 and 2016, respectively, with litterfall accounting for 60% of the total. Tidal forest soils in the streamside and the interior (0–60 cm) contained 3–6% organic C, 0.20–0.40% N, and 270–540 µg/g P. Soil accretion based on ¹³⁷Cs was 4.0 mm/year on the streamside and 0.2 mm/year in the forest interior. The rate of accretion in the interior is considerably less than the current rate of sea level rise (3.1 mm/year) along the Georgia coast. Because the accretion rate was much higher on the streamside, rates of C sequestration, N and P accumulation, and mineral sediment deposition also were much greater. Low accretion rates in the interior of the forest that accounts for most of the acreage suggests that accelerated sea level rise is likely to lead to foreseeable death of tidal forests from saltwater intrusion and submergence.

Despite the importance of tidal ecosystems in the global carbon budget, the relationships between environmental drivers and carbon dynamics in these wetlands remain poorly understood. This limited understanding results from the challenges associated with in situ flux studies and their correlation with satellite imagery which can be affected by periodic tidal flooding. Carbon dioxide eddy covariance (EC) towers are installed in only a few wetlands worldwide, and the longest eddy-covariance record from Georgia (GA) wetlands contains only two continuous years of observations. The goals of the present study were to evaluate the performance of existing MODIS Gross Primary Production (GPP) products (MOD17A2) against EC derived GPP and develop a tide-robust Normalized Difference Moisture Index (NDMI) based model to predict GPP within a Spartina alterniflora salt marsh on Sapelo Island, GA. These EC tower-based observations represent a basis to associate CO₂ fluxes with canopy reflectance and thus provide the means to use satellite-based reflectance data for broader scale investigations. We demonstrate that Light Use Efficiency (LUE)-based MOD17A2 does not accurately reflect tidal wetland GPP compared to a simple empirical vegetation index-based model where tidal influence was accounted for. The NDMI-based GPP model was capable of predicting changes in wetland CO₂ fluxes and explained 46% of the variation in flux-estimated GPP within the training data, and a root mean square error of 6.96 g C m⁻² in the validation data. Our investigation is the first to create a MODIS-based wetland GPP estimation procedure that demonstrates the importance of filtering tidal observations from satellite surface reflectance data.

Understanding which compounds comprising the complex and dynamic marine dissolved organic matter (DOM) pool are important in supporting heterotrophic bacterial production remains a major challenge. We eliminated sources of labile phytoplankton products, advected terrestrial material and photodegradation products to coastal microbial communities by enclosing water samples in situ for 24 h in the dark. Bacterial genes for which expression decreased between the beginning and end of the incubation and chemical formulae that were depleted over this same time frame were used as indicators of bioavailable compounds, an approach that avoids augmenting or modifying the natural DOM pool. Transport‐ and metabolism‐related genes whose relative expression decreased implicated osmolytes, carboxylic acids, fatty acids, sugars and organic sulfur compounds as candidate bioreactive molecules. FT‐ICR MS analysis of depleted molecular formulae implicated functional groups ~ 30–40 Da in size cleaved from semi‐polar components of DOM as bioreactive components. Both gene expression and FT‐ICR MS analyses indicated higher lability of compounds with sulfur and nitrogen heteroatoms. Untargeted methodologies able to integrate biological and chemical perspectives can be effective strategies for characterizing the labile microbial metabolites participating in carbon flux.

Predators can significantly affect prey by removing prey individuals and by changing prey behavior. The tradeoff between foraging behavior and predation risk may result in a trophic cascade that can have important effects on ecosystem processes. For herbivores that can feed both above‐ and belowground, it is likely that predation risk affects the location of feeding. We tested whether two species of predatory marsh crabs affected feeding behavior of the herbivorous crab, Sesarma reticulatum. We found that predatory crabs could kill or injure Sesarma and that Sesarma did less damage to its food plant Spartina alterniflora in the presence of the more dangerous predator. Sesarma prefers to feed on and grows better on belowground rhizomes than aboveground leaves; however, the costs of digging burrows to access rhizomes lead to higher mortality if it is the only diet option. The location of feeding did not affect total biomass of S. alterniflora. For Sesarma, a choice in feeding location allows the crabs the behavioral flexibility to balance the risks of predation, the nutritional benefit of feeding belowground, and the survival costs of belowground feeding. Similar tradeoffs are likely to increase the success of other herbivores that can feed both above‐ and belowground.