Publications

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).

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 photochemistryby 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.

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).

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.