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  1. Abstract We present new chronologies that inform the timing and tempo of shell ring and shell mound construction on the South Atlantic Bight. Our project combines recently acquired dates with legacy radiocarbon dates from over 25 rings and mounds to provide a higher-resolution chronology regarding the occupation and formation of this larger landscape of the earliest fishing villages along the East Coast of the United States. We resolve the ordering and timing of occupation of these rings and mounds through Bayesian statistical modeling. These new models historicize and contextualize these shell rings in ways previously impossible. Specifically, our new chronologies of these villages indicate that the earliest villages were established prior to the invention of pottery. The early period of village establishment evidences isolated village rings, whereas later periods seem to have more villages, but these appear to have been relocated to other areas and/or islands over time. Shell mounds are fewer in number, are spread throughout the time period, and may represent special purpose sites compared to shell-rings. Once villages spread, they quickly adopted new technologies (i.e., pottery) and created new institutions and practiced village relocation, which allowed this way of life to persist for more than a thousand years. 
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  2. Abstract Understanding how climate and local stressors interact is paramount for predicting future ecosystem structure. The effects of multiple stressors are often examined in small‐scale and short‐term field experiments, limiting understanding of the spatial and temporal generality of the findings. Using a 22‐year observational dataset of plant and grazer abundance in a southeastern US salt marsh, we analyzed how changes in drought and grazer density combined to affect plant biomass. We found: (1) increased drought severity and higher snail density both correlated with lower plant biomass; (2) drought and snail effects interacted additively; and, (3) snail effects had a threshold, with additive top‐down effects only occurring when snails were present at high densities. These results suggest that the emergence of multiple stressor effects can be density dependent, and they validate short‐term experimental evidence that consumers can augment environmental stress. These findings have important implications for predicting future ecosystem structure and managing natural ecosystems. 
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  3. Abstract Circular shell rings along the South Atlantic coast of the United States are vestiges of the earliest sedentary villages in North America, dating to 4500–3000 BP. However, little is known about when Indigenous communities began constructing these shell-ring villages. This article presents data from the Hokfv-Mocvse Shell Ring on Ossabaw Island, Georgia. Although shell rings are often associated with the earliest ceramics in North America, no ceramics were encountered in our excavations at Hokfv-Mocvse, and the only materials recovered were projectile points similar to points found over 300 km inland. Bayesian modeling of radiocarbon dates indicates that the ring was occupied between 5090 and 4735 cal BP (95% confidence), making it the earliest dated shell ring in the region. Additionally, shell geochemistry and oyster paleobiology data suggest that inhabitants were living at the ring year-round and had established institutions at that time to manage oyster fisheries sustainably. Hokfv-Mocvse therefore provides evidence for Indigenous people settling in year-round villages and adapting to coastal environments in the region centuries before the adoption of pottery. The establishment of villages marks a visible archaeological shift toward settling down and occupying island ecosystems on a more permanent basis and in larger numbers than ever before in the region. 
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  4. Abstract Predators regulate communities through top‐down control in many ecosystems. Because most studies of top‐down control last less than a year and focus on only a subset of the community, they may miss predator effects that manifest at longer timescales or across whole food webs. In southeastern US salt marshes, short‐term and small‐scale experiments indicate that nektonic predators (e.g., blue crab, fish, terrapins) facilitate the foundational grass,Spartina alterniflora, by consuming herbivorous snails and crabs. To test both how nekton affect marsh processes when the entire animal community is present, and how prior results scale over time, we conducted a 3‐year nekton exclusion experiment in a Georgia salt marsh using replicated 19.6 m2plots. Our nekton exclusions increased densities of plant‐grazing snails and juvenile deposit‐feeding fiddler crab and, in Year 2, reduced predation on tethered juvenile snails, indicating that nektonic predators control these key macroinvertebrates. However, in Year 3, densities of mesopredatory benthic mud crabs increased threefold in nekton exclusions, erasing the tethered snails' predation refuge. Nekton exclusion had no effect onSpartinabiomass, likely because the observed mesopredator release suppressed grazing snail densities and elevated densities of fiddler crabs, whose burrowing alleviates soil stresses. Structural equation modeling supported the hypotheses that nektonic predators and mesopredators control invertebrate communities, with nektonic predators having stronger total effects onSpartinathan mud crabs by controlling densities of species that both suppress (grazers) and facilitate (fiddler crabs) plant growth. These findings highlight that salt marshes can be resilient to multiyear reductions in nektonic predators if mesopredators are present and that multiple pathways of trophic control manifest in different ways over time to mediate community dynamics. These results highlight that larger scale and longer‐term experiments can illuminate community dynamics not previously understood, even in well‐studied ecosystems such as salt marshes. 
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  5. IntroductionDissolved organic matter (DOM) composition varies over space and time, with a multitude of factors driving the presence or absence of each compound found in the complex DOM mixture. Compounds ubiquitously present across a wide range of river systems (hereafter termed core compounds) may differ in chemical composition and reactivity from compounds present in only a few settings (hereafter termed satellite compounds). Here, we investigated the spatial patterns in DOM molecular formulae presence (occupancy) in surface water and sediments across 97 river corridors at a continental scale using the “Worldwide Hydrobiogeochemical Observation Network for Dynamic River Systems—WHONDRS” research consortium. MethodsWe used a novel data-driven approach to identify core and satellite compounds and compared their molecular properties identified with Fourier-transform ion cyclotron resonance mass spectrometry (FT-ICR MS). ResultsWe found that core compounds clustered around intermediate hydrogen/carbon and oxygen/carbon ratios across both sediment and surface water samples, whereas the satellite compounds varied widely in their elemental composition. Within surface water samples, core compounds were dominated by lignin-like formulae, whereas protein-like formulae dominated the core pool in sediment samples. In contrast, satellite molecular formulae were more evenly distributed between compound classes in both sediment and water molecules. Core compounds found in both sediment and water exhibited lower molecular mass, lower oxidation state, and a higher degree of aromaticity, and were inferred to be more persistent than global satellite compounds. Higher putative biochemical transformations were found in core than satellite compounds, suggesting that the core pool was more processed. DiscussionThe observed differences in chemical properties of core and satellite compounds point to potential differences in their sources and contribution to DOM processing in river corridors. Overall, our work points to the potential of data-driven approaches separating rare and common compounds to reduce some of the complexity inherent in studying riverine DOM. 
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  6. Abstract People often modify the shoreline to mitigate erosion and protect property from storm impacts. The 2 main approaches to modification are gray infrastructure (e.g., bulkheads and seawalls) and natural or green infrastructure (NI) (e.g., living shorelines). Gray infrastructure is still more often used for coastal protection than NI, despite having more detrimental effects on ecosystem parameters, such as biodiversity. We assessed the impact of gray infrastructure on biodiversity and whether the adoption of NI can mitigate its loss. We examined the literature to quantify the relationship of gray infrastructure and NI to biodiversity and developed a model with temporal geospatial data on ecosystem distribution and shoreline modification to project future shoreline modification for our study location, coastal Georgia (United States). We applied the literature‐derived empirical relationships of infrastructure effects on biodiversity to the shoreline modification projections to predict change in biodiversity under different NI versus gray infrastructure scenarios. For our study area, which is dominated by marshes and use of gray infrastructure, when just under half of all new coastal infrastructure was to be NI, previous losses of biodiversity from gray infrastructure could be mitigated by 2100 (net change of biodiversity of +0.14%, 95% confidence interval −0.10% to +0.39%). As biodiversity continues to decline from human impacts, it is increasingly imperative to minimize negative impacts when possible. We therefore suggest policy and the permitting process be changed to promote the adoption of NI. 
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  7. Abstract Large grazers modify vegetated ecosystems and are increasingly viewed as keystone species in trophic rewilding schemes. Yet, as their ecosystem influences are context‐dependent, a crucial challenge is identifying where grazers sustain, versus undermine, important ecosystem properties and their resilience.Previous work in diverse European saltmarshes found that, despite changing plant and invertebrate community structure, grazers do not suppress below‐ground properties, including soil organic carbon (SOC). We hypothesised that, in contrast, eastern US saltmarshes would be sensitive to large grazers as extensive areas are dominated by a single grass,Spartina alterniflora. We predicted that grazers would reduce above‐ and below‐groundSpartinabiomass, suppress invertebrate densities, shift soil texture and ultimately reduce SOC concentration.We tested our hypotheses using a replicated 51‐month large grazer (horse) exclusion experiment in Georgia, coupled with observations of 14 long‐term grazed sites, spanning ~1000 km of the eastern US coast.Grazer exclusion quickly led to increasedSpartinaheight, cover and flowering, and increased snail density. Changes in vegetation structure were reflected in modified soil texture (reduced sand, increased clay) and elevated root biomass, yet we found no response of SOC. Large grazer exclusion also reduced drought‐associated vegetation die‐off.We also observed vegetation shifts in sites along the eastern US seaboard where grazing has occurred for hundreds of years. Unlike in the exclusion experiment, long‐term grazing was associated with reduced SOC. A structural equation model implicated grazing by revealing reduced stem height as a key driver of reduced soil organic carbon.Synthesis: These results illustrate the context dependency of large grazer impacts on ecosystem properties in coastal wetlands. In contrast to well‐studied European marshes, eastern US marshes are dominated and structured by a single foundational grass species resulting in vegetation and soil properties being more sensitive to grazing. Coastal systems characterised by a single foundation species might be inherently vulnerable to large grazers and lack resilience in the face of other disturbances, underlining that frameworks to explain and predict large grazer impacts must account for geographic variation in ecosystem structure. 
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  8. Abstract Ecosystems vary broadly in their responses to disturbance, ranging from highly impacted to resilient or resistant. We conducted a large‐scale analysis of hurricane disturbance effects on coastal marshes by examining 20 years of data from 10 sites covering 100,000 ha at the Georgia Coastal Ecosystems Long‐Term Ecological Research site distributed across gradients of salinity and proximity to the ocean. We analyzed the impacts of Hurricanes Matthew (in 2016) and Irma (in 2017) on marsh biota (plants, crabs, and snails) and physical attributes (erosion, wrack deposition, and sedimentation). We compared these variables prior to the storms (2000–2015) to years with storms (2016, 2017) to those after the storms (2018–2020). Hurricanes generated storm surges that increased water depth and salinity of oligotrophic areas for up to 48 h. Biological variables in the marsh showed few effects of the hurricanes. The only physical variable affected was creek bank slumping; however, slumping had already increased a year before the hurricanes, suggesting that slumping could have a different cause. Thus, our study uncovered only minor, ephemeral impacts on Georgia coastal marshes, highlighting their resistance to hurricane disturbance of the lower magnitude that typically confronts this region of coastline. 
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  9. Abstract Tidal salt marshes are important ecosystems in the global carbon cycle. Understanding their net carbon exchange with the atmosphere is required to accurately estimate their net ecosystem carbon budget (NECB). In this study, we present the interannual net ecosystem exchange (NEE) of CO2derived from eddy covariance (EC) for aSpartina alterniflorasalt marsh. We found interannual NEE could vary up to 3‐fold and range from −58.5 ± 11.3 to −222.9 ± 12.4 g C m−2 year−1in 2016 and 2020, respectively. Further, we found that atmospheric CO2fluxes were spatially dependent and varied across short distances. High biomass regions along tidal creek and estuary edges had up to 2‐fold higher annual NEE than lower biomass marsh interiors. In addition to the spatial variation of NEE, regions of the marsh represented by distinct canopy zonation responded to environmental drivers differently. Low elevation edges (with taller canopies) had a higher correlation with river discharge (R2 = 0.61), the main freshwater input into the system, while marsh interiors (with short canopies) were better correlated with in situ precipitation (R2 = 0.53). Lastly, we extrapolated interannual NEE to the wider marsh system, demonstrating the potential underestimation of annual NEE when not considering spatially explicit rates of NEE. Our work provides a basis for further research to understand the temporal and spatial dynamics of productivity in coastal wetlands, ecosystems which are at the forefront of experiencing climate change induced variability in precipitation, temperature, and sea level rise that have the potential to alter ecosystem productivity. 
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  10. Abstract In salt marshes of the Southeastern USA, purple marsh crabs (Sesarma reticulatum), hereafterSesarma, aggregate in grazing and burrowing fronts at the heads of tidal creeks, accelerating creek incision into marsh platforms. We explored the effects of this keystone grazer and sediment engineer on salt marsh sediment accumulation, hydrology, and carbon (C) and nitrogen (N) turnover using radionuclides (210Pb and7Be), total hydrolyzable amino acids (THAA), and C and N stable isotopes (δ13C and δ15N) in sediment from pairedSesarma-grazed and un-grazed creeks.Sesarma-grazed-creek sediments exhibited greater bioturbation and tidal inundation compared to sediments in un-grazed creeks, as indicated by larger210Pb and7Be inventories. Total organic carbon (TOC) to total nitrogen (TN) weight ratios (C:N) were higher and δ15N values were lower in grazed-creek sediments than in un-grazed-creek sediments, suggestingSesarmaremove and assimilate N in their tissues, and excrete N with lower δ15N values into sediments. In support of this inference, the percent total carbon (TC) and percent TOC declined by nearly half, percent TN decreased by ~ 80%, and the C:N ratio exhibited a ~ threefold increase betweenSesarmafore-gut and hind-gut contents. An estimated 91% ofSesarma’s diet was derived fromSpartina alterniflora,the region’s dominant salt marsh plant. We found that, asSesarmagrazing fronts progress across marsh landscapes, they enhance the decay ofSpartina-derived organic matter and prolong marsh tidal inundation. These findings highlight the need to better account for the effects of keystone grazers and sediment engineers, likeSesarma, in estimates of the stability and size of blue C stores in coastal wetlands. 
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