Recent Publications
Jung, Y. and Burd, A.B. 2017. Seasonal changes in above- and below-ground non-structural carbohydrates (NSC) in Spartina alterniflora in a marsh in Georgia, USA. Aquatic Botany. 140:13-22. (DOI: https://doi.org/10.1016/j.aquabot.2017.04.003)
Abstract
Spartina alterniflora is the dominant grass appearing in salt marshes along the east coast of the USA. The development of predictive, mechanistic models of Spartina has been hindered by the lack of information on below-ground biomass and its dynamics, and in particular the storage of resources that can be used for spring re-growth. We studied the dynamics of non-structural carbohydrates (glucose, fructose, sucrose, and starch) and biomass in 8 different above- and below-ground tissues in S. alternifloraover the course of a year in a salt marsh on Sapelo Island, Georgia, USA. We found greater seasonal variability in non-structural carbohydrates in S. alterniflora than had been previously reported, with concentrations varying between 3.3% through 17.3% of the total biomass and between 0% and 19.5% of dry weight depending on the type of tissue, with statistical differences between the different tissues. We found that sucrose was the dominant non-structural carbohydrate in above- and below-ground tissue, and that this sugar was likely used for long-term storage during winter months and as a resource for early spring growth. Glucose, fructose, and starch showed less variability, with glucose following changes in above-ground biomass more closely indicating their use as short-term storage. We were unable to develop a coherent carbon budget for the plants largely because of uncertainties in modeled net primary production and heterogeneity in below-ground biomass.
Abstract
In salt marshes, disturbance by wrack (floating mats of dead vegetation) is common and affects plant productivity and species composition, but little is known about how the timing of disturbance mediates these effects, nor how it interacts with herbivory. Using a field experiment on the Georgia coast, we simulated the effects of wrack disturbance at different times of the year on the marsh grass Spartina alterniflora and its stem-boring herbivorous insects. The timing of disturbance throughout the growing season strongly affected fall biomass, stem height, the proportion of stems flowering, and the proportion of stems colonized by stem-boring herbivorous insects. End-of-season biomass in plots disturbed in March did not differ from undisturbed controls, but biomass was reduced by 50% in plots disturbed in May, and by over 90% in plots disturbed in September. Disturbance in March and May stimulated flowering, but disturbance later in the growing season suppressed it. Plots disturbed late in the growing season had a low frequency of stem-boring herbivores. Stems containing stem borers rarely flowered. These results indicate that the timing of disturbance matters in coastal salt marshes. Late-season disturbances had the strongest effects on S. alterniflora and its herbivores. Disturbances early in the growing season did not affect end-of-season biomass, and stimulated flowering, suggesting parallels between fire disturbance in grasslands and wrack disturbance in salt marshes. Late-season disturbance did reduce herbivory by stem-boring insects, but not enough to compensate for the direct effects of disturbance on the plants. Future studies of disturbance in salt marshes should consider how the timing of experimental disturbance treatments relates to the timing of natural disturbances.
Abstract
Microbially-mediated transformations of dissolved organic matter (DOM) in a marsh-dominated estuarine system were investigated at the molecular level using ultrahigh resolution mass spectrometry. In addition to observing spatial and temporal variability in DOM sources in the estuary, multiple incubations with endogenous microorganisms identified the influence of DOM composition on biodegradation. A clear microbial preference for degradation of compounds associated with marine DOM relative to those of terrestrial origin was observed, resulting in an overall shift of the remaining DOM toward a stronger terrigenous signature. During short, 1-day long incubations of samples rich in marine DOM, the molecular formulae that were enriched had slightly smaller mass (20–30 Da) and number of carbon atoms compared to the molecular formulae that were depleted. Over longer time scales (70 days), the mean differences in molecular mass between formulae that were depleted and enriched were substantially larger (~270 Da). The differences in elemental composition over daily time scales were consistent with transformations in functional groups; over longer time scales, the differences in elemental composition may be related to progressive transformations of functional groups of intermediate products and/or other reactions. Our results infused new data toward the understanding of DOM processing by bacterioplankton in estuarine systems.
Abstract
Ecosystem engineers affect different organisms and processes in multiple ways at different spatial scales. Moreover, similar species may differ in their engineering effects for reasons that are not always clear. We examined the role of four species of burrowing crabs (Sesarma reticulatum, Eurytium limosum, Panopeus herbstii, Uca pugnax) in engineering tidal creek networks in salt marshes experiencing sea level rise. In the field, crab burrows were associated with heads of eroding creeks and the loss of plant (Spartina alterniflora) stems. S. reticulatum was closely associated with creek heads, but densities of the other crab species did not vary across marsh zones. In mesocosm experiments, S. reticulatum excavated the most soil and strongly reduced S. alterniflora biomass. The other three species excavated less and did not affect S. alterniflora. Creek heads with vegetation removed to simulate crab herbivory grew significantly faster than controls. Percolation rates of water into marsh sediments were 10 times faster at creek heads than on the marsh platform. Biomass decomposed two times faster at creek heads than on the marsh platform. Our results indicate that S. reticulatum increases creek growth by excavating sediments and by consuming plants, thereby increasing water flow and erosion at creek heads. Moreover, it is possible that S. reticulatum burrows also increase creek growth by increasing surface and subsurface erosion, and by increasing decomposition of organic matter at creek heads. Our results show that the interaction between crab and plant ecosystem engineers can have both positive and negative effects. At a small scale, in contrast to other marsh crabs, S. reticulatum harms rather than benefits plants, and increases erosion rather than marsh growth. At a large scale, however, S. reticulatum facilitates the drainage efficiency of the marsh through the expansion of tidal creek networks, and promotes marsh health.
Abstract
The concentrations of dissolved copper (Cud), copper-binding ligands, thiourea-type thiols, and humic substances (HSCu) were measured in estuarine waters adjacent to Sapelo Island, Georgia, USA, on a monthly basis from April to December 2014. Here we present the seasonal cycle of copper speciation within the estuary and compare it to the development of an annually occurring bloom of Ammonia Oxidizing Archaea (AOA), which require copper for many enzymes. Two types of complexing ligands (L1 and L2) were found to dominate with mean complex stabilities (log K′CuLKCuL′) of 14.5 and 12.8. Strong complexation resulted in lowering the concentration of free cupric ion (Cu2+) to femtomolar (fM) levels throughout the study and to sub-fM levels during the summer months. A Thaumarchaeota bloom during this period suggests that this organism manages to grow at very low Cu2+ concentrations. Correlation of the concentration of the L1 ligand class with a thiourea-type thiol and the L2 ligand class with HSCuprovide an interesting dimension to the identity of the ligand classes. Due to the stronger complex stability, 82–99% of the copper was bound to L1. Thiourea-type thiols typically form Cu(I) species, which would suggest that up to ~90% copper could be present as Cu(I) in this region. In view of the very low concentration of free copper (pCu > 15 at the onset and during the bloom) and a reputedly high requirement for copper, it is likely that the Thaumarchaeota are able to access thiol-bound copper directly.
Abstract
Oysters provide a critical habitat, are a food resource for higher trophic levels and support important commercial fisheries throughout the world. Oyster reefs can improve water quality by removing phytoplankton. While sediment denitrification may be enhanced adjacent to oyster reefs, little is known about nitrification and denitrification associated with living oysters and their shells. We measured nitrification and denitrification in living oysters (Crassostrea virginica and Crassostrea gigas) and empty oyster shells. Nitrification was similar between live oysters and empty oyster shells, however, denitrification was enhanced significantly on living oysters compared to shells. This is the first demonstration of nitrification and denitrification associated with living oysters and their shells. Our data suggest that loss of historic oyster reefs has likely affected the resilience of estuaries to
eutrophication. The additional benefit of oyster mediated denitrification should be considered in restoration of oyster reefs as a tool for managing eutrophication.
Abstract
We measured sediment organic carbon and nitrogen accumulation and rates of denitrification enzyme activity and greenhouse gas (CO2, CH4, N2O) production from slurries of sediments of a mudflat that formed in 2002, a young (8-year-old) natural Spartina alterniflora salt marsh that developed on part of the mudflat, and four mature (>200 years old) salt marshes in southeastern Georgia to examine microbial processes related to carbon (C) and nitrogen (N) cycling during succession from mudflat to mature marsh. Soil organic C and N and C: N ratio (0–30 cm) increased across the chronosequence from mudflat (791 ± 35 g C/m2, 125 ± 17 g N/m2) to young marsh (2520 ± 131 g C/m2, 190 ± 10 g N/m2) to mature marshes (5827 ± 250 g C/m2, 372 ± 20 g N/m2). After 8 years of colonization by S. alterniflora, sediment organic carbon increased 3.2 times, and nitrogen increased 1.5 times relative to the mudflat. The high rate of organic C and N accumulation based on time series measurements (188 g C/m2/yr, 7.8 g N/m2/yr) and feldspar marker layers (359 g C/m2/yr, 26.2 g N/m2/yr) was attributed to high accretion (3 cm/yr) in this low elevation (0.18 m NAVD88) emerging marsh. Carbon dioxide production increased with increasing sediment organic C from mudflat to mature marshes. Un-amended denitrification enzyme activity, measured in slurry incubations, ranged from an average of 0.020 ± 0.005 μg g−1 hr−1 in the mature marshes to 0.094 ± 0.03 μg g−1hr−1 in the young marsh. We also measured denitrification potential in slurry incubations amended with C (glucose), N (nitrate), and C + N to assess the potential for substrate limitations. Denitrification potential in the mudflat did not show strong nutrient limitation. In the young marsh, denitrification potential was C-limited, and in the mature marsh, it was co-limited by C and N. In July samples, CO2 production showed a statistically significant increase with age from the mudflat to the mature marshes. However, in both months, CO2 production efficiency (expressed on a per g C basis) was significantly higher in the mudflat sediment slurries than in the young marsh and mature marsh samples. Spartina colonization of mudflats and the subsequent accumulation of organic matter are key to enriching sediment organic C and N pools that control microbial heterotrophy, particularly denitrification and CO2production, which play important roles in marsh C and N cycling.
Abstract
Disturbance regimes are shifting in response to climate change, land-use change, species’ invasions, and other stressors, challenging ecologists to improve understanding of the mechanisms controlling plant recovery under different conditions. In this study, we investigate mechanisms that underpin plant recolonization of two types of disturbance: disturbances that remove standing plant biomass, but leave the underlying soil structure largely intact, and those that remove standing biomass and physically disrupt soil structure. In the southeastern United States, salt marshes, drought and invasive feral hogs (Sus scrofa) are associated with disturbances of these respective types and both leave behind mudflats dotted with patches of cordgrass (Spartina alterniflora), the system’s dominant foundation species. To test how disturbance type and remnant patch size may interact to affect cordgrass recolonization, we transplanted replicate cordgrass patches of three sizes into one mudflat that experienced vegetation-only disturbance (VD) during a recent drought, and a second mudflat where hog activities had disrupted the soil structure creating a soil + vegetation disturbance (soil + VD). Over one year, we monitored plant performance and properties of soil. Compared with the drought-associated VD where patches, regardless of their size, expanded vigorously, large and medium patches grew little and smaller patches even less in the hog-associated soil + VD mudflat. Moreover while biogeochemical properties varied little in the VD compared with adjacent vegetated marsh areas, the mudflats with soil + VD had less soil-binding organic matter and, hence, were too soft to support crab burrows, leading to a reduction in oxygen availability and cordgrass expansion. These results indicate that cordgrass recovery is far faster from disturbances that do not degrade soil structure than those that do and therefore advocate for disturbance-specific management strategies. Specifically, while transplanting patches is effective in restoring marshes disturbed by drought, wrack, or other factors that leave soil structure intact, preventative measures, like hog population culling, are essential to mitigating the ecological impacts of soil structure-altering disturbances.
Tolar, B., Powers, L.C., Miller, W., Wallsgrove, N.J., Popp, B.N. and Hollibaugh, J.T. 2016. Ammonia oxidation in the ocean can be inhibited by nanomolar concentrations of hydrogen peroxide. Frontiers in Marine Science,| https://doi.org/10.3389/fmars.2016.00237.
Abstract
Marine Thaumarchaeota were discovered over 20 years ago and although a few isolates from this group are now available for study, we do not yet understand the environmental controls on their growth and distribution. Thaumarchaeotes oxidize ammonia to nitrite, mediating a key step in the global nitrogen cycle, and it is estimated that about 20% of all prokaryotic cells in the ocean belong to this phylum. Despite their almost ubiquitous distribution, marine Thaumarchaeota are rarely abundant in open-ocean surface (<100 m) waters. We tested the hypothesis that this vertical distribution is driven by reactive oxygen species (ROS), specifically H2O2, generated by photochemical and biological processes—“indirect photoinhibition” rather than light inhibition as previously postulated for ammonia-oxidizing Archaea. Here we show that H2O2 can be surprisingly toxic to Thaumarchaeota from the Southern Ocean, with ammonia oxidation inhibited by additions of as little as 10 nM H2O2, while temperate Thaumarchaeota ecotypes were more tolerant. This sensitivity could explain the seasonal disappearance of Thaumarchaeota from polar surface waters and the increase in ammonia oxidation rates with depth commonly observed in marine environments. Our results highlight the need for further physiological studies of Thaumarchaeota, and indicate that ROS sensitivity could be used as a characteristic for dividing the group into meaningful ecotypes.
Abstract
Urea nitrogen has been proposed to contribute significantly to nitrification by marine thaumarchaeotes. These inferences are based on distributions of thaumarchaeote urease genes rather than activity measurements. We found that ammonia oxidation rates were always higher than oxidation rates of urea-derived N in samples from coastal Georgia, USA (means 6 SEM: 382 6 35 versus 73 6 24 nmol L21 d21 , Mann-Whitney U-test p < 0.0001), and the South Atlantic Bight (20 6 8.8 versus 2.2 6 1.7 nmol L21 d21 , p 5 0.026) but not the Gulf of Alaska (8.8 6 4.0 versus 1.5 6 0.6, p > 0.05). Urea-derived N was relatively more important in samples from Antarctic continental shelf waters, though the difference was not statistically significant (19.4 6 4.8 versus 12.0 6 2.7 nmol L21 d21 , p > 0.05). We found only weak correlations between oxidation rates of urea-derived N and the abundance or transcription of putative Thaumarchaeota ureC genes. Dependence on urea-derived N does not appear to be directly related to pH or ammonium concentrations. Competition experiments and release of 15NH3 suggest that urea is hydrolyzed to ammonia intracellularly, then a portion is lost to the dissolved pool. The contribution of urea-derived N to nitrification appears to be minor in temperate coastal waters, but may represent a significant portion of the nitrification flux in Antarctic coastal waters.
Abstract
A high-resolution coastal ocean model is used to investigate salinity variability and water exchange in a complex coastal system off the southern U.S. characterized by three adjacent sounds that are interconnected by a network of channels, creeks, and intertidal areas. Model results are generally highly correlated with observations from the Georgia Coastal Ecosystem Long Term Ecological Research (GCE-LTER) program, revealing a high degree of salinity variability at the Altamaha River and Doboy Sound, decreasing sharply toward Sapelo Sound. A Lagrangian particle tracking method is used to investigate local residence time and connectivity in the system. Local residence time is highly variable, increasing with distance from the Altamaha River and decreasing with river flow, revealing that discharge plays a dominant role on transport processes and estuary-shelf exchange. The Altamaha River and Doboy Sound are connected to each other in all seasons, with exchange occurring both via coastal and estuarine pathways. While particles released at the Altamaha and Doboy rarely reach Sapelo Sound, particles released at Sapelo Sound and the creeks surrounding its main channel can reach the entire estuarine system.
Abstract
The establishment of foundation plants in bare mudflats is a critical process. While consumers are increasingly recognized to exert strong top-down control of plant performance in salt marshes, studies to date have focused on the effects of consumers on mature stands rather than on plants that are recolonizing after disturbance or where restoration has occurred. Furthermore, whether consumer-facilitated fungal infection differentially affects newly establishing plants in mudflats compared to mature stands remains poorly understood. In a salt marsh in southern Brazil, we examined the effects of herbivory by the crab Neohelice granulata and fungal infection on the survival and growth of Spartina alterniflora transplanted into mudflats. We additionally tested the effects of herbivory and fungi on newly established versus well-established stands of S. alterniflora. Highly intensive natural crab herbivory significantly reduced the development of S. alterniflora and increased its fungal infection by 50%. Light herbivory, removing only small areas of plant leaves, reduced the height growth and leaf production of directly affected tillers by about 14 to 18%, and both newly and well-established, clonally integrated stands of S. alterniflora allocated energy towards the formation of new tillers. While herbivory facilitated fungal infection and subsequent fungal damage in leaves, no significant effects of fungicide treatment or its interactions with crab grazing on S. alterniflora growth were detected, suggesting a saprophytic rather than a pathogenic role of fungi in this 3-species interaction. Here, we found that marsh grasses transplanted for restoration or those colonizing disturbance-generated mudflats may be facilitated by protection against consumers.
Abstract
We documented the frequency and effect on live biomass of five different types of disturbance over 14 years in creekbank and mid-marsh zones of eight salt marshes dominated by Spartina alterniflora in Georgia, USA. Wrack (floating debris) and creekbank slumping were the most common disturbances at the creekbank, and snails were the most common disturbance agent in the mid-marsh. Disturbance frequency varied among sites due to differences in plot elevation and landscape position. Wrack disturbance at the creekbank was positively correlated with plot elevation, and both initial slumping and terminal slumping of creekbank plots were negatively correlated with plot elevation. Wrack disturbance at the creekbank and snail disturbance in the mid-marsh were also most common at barrier island vs. interior marshes. Disturbance varied up to 14-fold among years. Wrack disturbance at the creekbank was negatively correlated with river discharge and sea level, and initial slumping of creekbank plots was also negatively correlated with sea level. The different disturbance types varied in their effects on end-of-year standing plant biomass. At the creekbank, wrack disturbance reduced biomass in affected plots by ~46%, but slumping did not affect biomass until the plot was totally lost. In the mid-marsh, slumping and wrack were not important disturbances, but snail disturbance reduced biomass in affected plots by ~70%. In addition, abiotic conditions (river discharge, maximum monthly temperature, sea level, and precipitation) strongly affected year-to-year variation in biomass. Across the entire landscape, fewer than a quarter of the plots on average were disturbed, and disturbance reduced overall standing biomass by ~18% in the creekbank zone and ~3% in the mid-marsh zone. Our results indicate that wrack has fairly strong effects on end-of-year biomass at the creekbank. Overall, however, variation in abiotic conditions among years had stronger effects on end-of-year standing biomass in both marsh zones than did disturbance.
Abstract
PhenoCams are part of a national network of automated digital cameras used to assess vegetation phenology transitions. Effectively analyzing PhenoCam time-series involves eliminating scenes with poor solar illumination or high cover of non-target objects such as water. We created a smart classifier to process images from the “GCESapelo” PhenoCam, which photographs a regularly-flooded salt marsh. The smart classifier, written in R, assigns pixels to target (vegetation) and non-target (water, shadows, fog and clouds) classes, allowing automated identification of optimal scenes for evaluating phenology. When compared to hand-classified validation images, the smart classifier identified scenes with optimal vegetation cover with 96% accuracy and other object classes with accuracies ranging from 86 to 100%. Accuracy for estimating object percent cover ranged from 74 to 100%. Pixel-classification with the smart classifier outperformed previous approaches (i.e. indices based on average color content within ROIs) and reduced variance in phenology index time-series. It can be readily adapted for other applications.
Abstract
Rapid, global, anthropogenic alteration of food webs in ecosystems necessitates a better understanding of how consumers regulate natural communities. We provide a global synthesis of consumer control of vegetation in coastal wetlands, where the domineering role of physical factors such as nutrient and salinity, rather than consumers, has been emphasized for decades. Using a data set of 1748 measures of consumer effects reported in 443 experiments/observations on all continents except Antarctica, we examine the generality of consumer control in salt marshes and mangroves globally. Our analyses show that salt marsh herbivores, including insects, snails, crabs, waterfowl, small mammals, and livestock, generally and often strongly suppress plant survival, aboveground biomass, and height, while their effects on plant density, belowground biomass, reproduction, and cover are more variable. These effects occur in forbs, grasses, and shrubs, and in both seedlings and adult plants. Herbivores additionally affect plant nutrient stoichiometry, and mediate plant interactions, though their effects on plant diversity are less consistent. Higher trophic levels also affect plants, as predators facilitate plant growth through trophic cascades that suppress grazer abundance and grazing rate. In mangroves, there are also signs of consumer control, though the relatively few studies available have often focused on mangrove propagules and seedlings rather than adults. Our analyses further reveal that the strength of consumer control is regulated by many physical factors. Nutrient, disturbance, and flooding, for example, amplify the negative effects of herbivores. Along latitudinal gradients, increased temperature enhances the negative effects of ectothermic herbivores, but has no effect on those of endothermic herbivores. Consumer control of coastal plants is also apparent across study methodologies: in field and laboratory settings, in observational studies, in consumer exclusion and addition experiments, in natural and transplanted plants, and in experiments of various durations. The role of consumer control in coastal vegetation worldwide highlights the need to better recognize and theoretically conceptualize both top-down and bottom-up forcing and their interactions in coastal wetlands. Improved understanding and conservation of coastal wetlands will only occur if we incorporate what the science has revealed: trophic feedbacks are an important and pervasive determinant of coastal plant communities.
Summary
- Phenotypic variation plays an important role in successful plant invasions. The spread of invasive species over large geographic ranges may be facilitated if plants can match their phenotype to local abiotic conditions. Spartina alterniflora, native to the United States, was introduced into China in 1979 and has spread over 19ᵒ of latitude along the eastern coast of China.
- We studied patterns in vegetative growth and sexual reproduction of S. alterniflora at 22 sites at 11 geographic locations over a latitudinal gradient of ˜2000 km from Tanggu (39.05°N, high latitude) to Leizhou (20.90°N, low latitude) in China. We further evaluated the basis of phenotypic differences by growing plants from across the range in a common garden for 2 growing seasons.
- We found distinct latitudinal clines in plant height, shoot density and sexual reproduction across latitude. Some traits exhibited linear relationships with latitude; others exhibited hump-shaped relationships. We identified correlations between plant traits and abiotic conditions such as mean annual temperature, growing degree days, tidal range and soil nitrogen content. However, geographic variation in all but one trait disappeared in the common garden, indicating that variation largely due to phenotypic plasticity. Only a slight tendency for latitudinal variation in seed set persisted for 2 years in the common garden, suggesting that plants may be evolving genetic clines for this trait.
- Synthesis. The rapid spread of Spartina alterniflora (S. alterniflora) in China has probably been facilitated by phenotypic plasticity in growth and reproductive traits. We found little evidence for the evolution of genetic clines in China, even though these exist for some traits in the native range. The considerable variation among clones, within provenances, that persisted in the common garden suggests a potential for the evolution of geographic clines in the future. Low fecundity of low-latitude S. alterniflora populations in China might result in a slower spread at low latitudes, but S. alterniflora is likely to continue to spread rapidly at high latitudes in China and into the Korean peninsula.
Abstract
Droughts are increasing in severity and frequency, yet the mechanisms that strengthen ecosystem resilience to this stress remain poorly understood. Here, we test whether positive interactions in the form of a mutualism between mussels and dominant cordgrass in salt marshes enhance ecosystem resistance to and recovery from drought. Surveys spanning 250 km of southeastern US coastline reveal spatially dispersed mussel mounds increased cordgrass survival during severe drought by 5- to 25-times. Surveys and mussel addition experiments indicate this positive effect of mussels on cordgrass was due to mounds enhancing water storage and reducing soil salinity stress. Observations and models then demonstrate that surviving cordgrass patches associated with mussels function as nuclei for vegetative re-growth and, despite covering only 0.1–12% of die-offs, markedly shorten marsh recovery periods. These results indicate that mutualisms, in supporting stress-resistant patches, can play a disproportionately large, keystone role in enhancing ecosystem resilience to climatic extremes.
Abstract
We examined the influence of abiotic drivers on inter-annual and phenological patterns of aboveground biomass for Marsh Cordgrass, Spartina alterniflora, on the Central Georgia Coast. The linkages between drivers and plant response via soil edaphic factors are captured in our graphical conceptual model. We used geospatial techniques to scale up in situ measurements of aboveground S. alterniflora biomass to landscape level estimates using 294 Landsat 5 TM scenes acquired between 1984 and 2011. For each scene we extracted data from the same 63 sampling polygons, containing 1222 pixels covering about 1.1 million m2. Using univariate and multiple regression tests, we compared Landsat derived biomass estimates for three S. alterniflora size classes against a suite of abiotic drivers. River discharge, total precipitation, minimum temperature, and mean sea level had positive relationships with and best explained biomass for all dates. Additional results, using seasonally binned data, indicated biomass was responsive to changing combinations of variables across the seasons. Our 28-year analysis revealed aboveground biomass declines of 33%, 35%, and 39% for S. alterniflora tall, medium, and short size classes, respectively. This decline correlated with drought frequency and severity trends and coincided with marsh die-backs events and increased snail herbivory in the second half of the study period.