Year-around survey and manipulation experiments reveal differential sensitivities of soil prokaryotic and fungal communities to saltwater intrusion in Florida Everglades wetlands.

Global sea-level rise is transforming coastal ecosystems, especially freshwater wetlands, in part due to increased episodic or chronic saltwater exposure, leading to shifts in biogeochemistry, plant- and microbial communities, as well as ecological services. Yet, it is still difficult to predict how soil microbial communities respond to the saltwater exposure because of poorly understood microbial sensitivity within complex wetland soil microbial communities, as well as the high spatial and temporal heterogeneity of wetland soils and saltwater exposure. To address this, we first conducted a two-year survey of microbial community structure and bottom water chemistry in submerged surface soils from 14 wetland sites across the Florida Everglades. We identified ecosystem-specific microbial biomarker taxa primarily associated with variation in salinity. Bacterial, archaeal and fungal community composition differed between freshwater, mangrove, and marine seagrass meadow sites, irrespective of soil type or season. Especially, methanogens, putative denitrifying methanotrophs and sulfate reducers shifted in relative abundance and/or composition between wetland types. Methanogens and putative denitrifying methanotrophs declined in relative abundance from freshwater to marine wetlands, whereas sulfate reducers showed the opposite trend. A four-year experimental simulation of saltwater intrusion in a pristine freshwater site and a previously saltwater-impacted site corroborated the highest sensitivity and relative increase of sulfate reducers, as well as taxon-specific sensitivity of methanogens, in response to continuously pulsing of saltwater treatment. Collectively, these results suggest that besides increased salinity, saltwater-mediated increased sulfate availability leads to displacement of methanogens by sulfate reducers even at low or temporal salt exposure. These changes of microbial composition could affect organic matter degradation pathways in coastal freshwater wetlands exposed to sea-level rise, with potential consequences, such as loss of stored soil organic carbon.

Modeling net ecosystem carbon balance and loss in coastal wetlands exposed to sea-level rise and saltwater intrusion.

Coastal wetlands are globally important stores of carbon (C). However, accelerated sea-level rise (SLR), increased saltwater intrusion, and modified freshwater discharge can contribute to the collapse of peat marshes, converting coastal peatlands into open water. Applying results from multiple experiments from sawgrass (Cladium jamaicense)-dominated freshwater and brackish water marshes in the Florida Coastal Everglades, we developed a system-level mechanistic peat elevation model (EvPEM). We applied the model to simulate net ecosystem C balance (NECB) and peat elevation in response to elevated salinity under inundation and drought exposure. Using a mass C balance approach, we estimated net gain in C and corresponding export of aquatic fluxes ( F AQ $$ {F}_{\mathrm{AQ}} $$ ) in the freshwater marsh under ambient conditions (NECB = 1119 ± 229 gC m-2  year-1 ; FAQ  = 317 ± 186 gC m-2  year-1 ). In contrast, the brackish water marsh exhibited substantial peat loss and aquatic C export with ambient (NECB = -366 ± 15 gC m-2  year-1 ; FAQ  = 311 ± 30 gC m-2  year-1 ) and elevated salinity (NECB = -594 ± 94 gC m-2  year-1 ; FAQ  = 729 ± 142 gC m-2  year-1 ) under extended exposed conditions. Further, mass balance suggests a considerable decline in soil C and corresponding elevation loss with elevated salinity and seasonal dry-down. Applying EvPEM, we developed critical marsh net primary productivity (NPP) thresholds as a function of salinity to simulate accumulating, steady-state, and collapsing peat elevations. The optimization showed that ~150-1070 gC m-2  year-1 NPP could support a stable peat elevation (elevation change ≈ SLR), with the corresponding salinity ranging from 1 to 20 ppt under increasing inundation levels. The C budgeting and modeling illustrate the impacts of saltwater intrusion, inundation, and seasonal dry-down and reduce uncertainties in understanding the fate of coastal peat wetlands with SLR and freshwater restoration. The modeling results provide management targets for hydrologic restoration based on the ecological conditions needed to reduce the vulnerability of the Everglades' peat marshes to collapse. The approach can be extended to other coastal peatlands to quantify C loss and improve understanding of the influence of the biological controls on wetland C storage changes for coastal management.

Water levels primarily drive variation in photosynthesis and nutrient use of scrub Red Mangroves in the southeastern Florida Everglades.

We investigated how mangrove-island micro-elevation (i.e., habitat: center vs edge) affects tree physiology in a scrub mangrove forest of the southeastern Everglades. We measured leaf gas exchange rates of scrub Rhizophora mangle L. trees monthly during 2019, hypothesizing that CO2 assimilation (Anet) and stomatal conductance (gsw) would decline with increasing water levels and salinity, expecting more considerable differences at mangrove-island edges than centers, where physiological stress is greatest. Water levels varied between 0 and 60 cm from the soil surface, rising during the wet season (May-October) relative to the dry season (November-April). Porewater salinity ranged from 15 to 30 p.p.t., being higher at mangrove-island edges than centers. Anet maximized at 15.1 µmol m-2 s-1, and gsw was typically <0.2 mol m-2 s-1, both of which were greater in the dry than the wet season and greater at island centers than edges, with seasonal variability being roughly equal to variation between habitats. After accounting for season and habitat, water level positively affected Anet in both seasons but did not affect gsw. Our findings suggest that inundation stress (i.e., water level) is the primary driver of variation in leaf gas exchange rates of scrub mangroves in the Florida Everglades, while also constraining Anet more than gsw. The interaction between inundation stress due to permanent flooding and habitat varies with season as physiological stress is alleviated at higher-elevation mangrove-island center habitats during the dry season. Freshwater inflows during the wet season increase water levels and inundation stress at higher-elevation mangrove-island centers, but also potentially alleviate salt and sulfide stress in soils. Thus, habitat heterogeneity leads to differences in nutrient and water acquisition and use between trees growing in island centers versus edges, creating distinct physiological controls on photosynthesis, which likely affect carbon flux dynamics of scrub mangroves in the Everglades.

Hurricanes fertilize mangrove forests in the Gulf of Mexico (Florida Everglades, USA).

Hurricanes are recurring high-energy disturbances in coastal regions that change community structure and function of mangrove wetlands. However, most of the studies assessing hurricane impacts on mangroves have focused on negative effects without considering the positive influence of hurricane-induced sediment deposition and associated nutrient fertilization on mangrove productivity and resilience. Here, we quantified how Hurricane Irma influenced soil nutrient pools, vertical accretion, and plant phosphorus (P) uptake after its passage across the Florida Coastal Everglades in September 2017. Vertical accretion from Irma's deposits was 6.7 to 14.4 times greater than the long-term (100 y) annual accretion rate (0.27 ± 0.04 cm y-1). Storm deposits extended up to 10-km inland from the Gulf of Mexico. Total P (TP) inputs were highest at the mouth of estuaries, with P concentration double that of underlying surface (top 10 cm) soils (0.19 ± 0.02 mg cm-3). This P deposition contributed 49 to 98% to the soil nutrient pool. As a result, all mangrove species showed a significant increase in litter foliar TP and soil porewater inorganic P concentrations in early 2018, 3 mo after Irma's impact, thus underscoring the interspecies differences in nutrient uptake. Mean TP loading rates were five times greater in southwestern (94 ± 13 kg ha-1 d-1) mangrove-dominated estuaries compared to the southeastern region, highlighting the positive role of hurricanes as a natural fertilization mechanism influencing forest productivity. P-rich, mineral sediments deposited by hurricanes create legacies that facilitate rapid forest recovery, stimulation of peat soil development, and resilience to sea-level rise.

Disturbance legacies increase and synchronize nutrient concentrations and bacterial productivity in coastal ecosystems.

Long-term ecological research can resolve effects of disturbance on ecosystem dynamics by capturing the scale of disturbance and interactions with environmental changes. To quantify how disturbances interact with long-term directional changes (sea-level rise, freshwater restoration), we studied 17 yr of monthly dissolved organic carbon (DOC), total nitrogen (TN), and phosphorus (TP) concentrations and bacterioplankton productivity across freshwater-to-marine estuary gradients exposed to multiple disturbance events (e.g., droughts, fire, hurricanes, and low-temperature anomalies) and long-term increases in water levels. By studying two neighboring drainages that differ in hydrologic connectivity, we additionally tested how disturbance legacies are shaped by hydrologic connectivity. We predicted that disturbance events would interact with long-term increases in water levels in freshwater and marine ecosystems to increase spatiotemporal similarity (i.e., synchrony) of organic matter, nutrients, and microbial activities. Wetlands along the larger, deeper, and tidally influenced Shark River Slough (SRS) drainage had higher and more variable DOC, TN, and TP concentrations than wetlands along the smaller, shallower, tidally restricted Taylor River Slough/Panhandle (TS/Ph) drainage. Along SRS, DOC concentrations declined with proximity to coast, and increased in magnitude and variability following drought and flooding in 2015 and a hurricane in 2017. Along TS/Ph, DOC concentrations varied by site (higher in marine than freshwater wetlands) but not year. In both drainages, increases in TN from upstream freshwater marshes occurred following fire in 2008 and droughts in 2010 and 2015, whereas downstream increases in TP occurred with coastal storm surge from hurricanes in 2005 and 2017. Decreases in DOC:TN and DOC:TP were explained by increased TN and TP. Increases in bacterioplankton productivity occurred throughout both drainages following low-temperature events (2010 and 2011) and a hurricane (2017). Long-term TN and TP concentrations and bacterioplankton productivity were correlated (r > 0.5) across a range of sampling distances (1-50 km), indicating spatiotemporal synchrony. DOC concentrations were not synchronized across space or time. Our study advances disturbance ecology theory by illustrating how disturbance events interact with long-term environmental changes and hydrologic connectivity to determine the magnitude and extent of disturbance legacies. Understanding disturbance legacies will enhance prediction and enable more effective management of rapidly changing ecosystems.

Phosphorus alleviation of salinity stress: effects of saltwater intrusion on an Everglades freshwater peat marsh.

Saltwater intrusion and salinization of coastal wetlands around the world are becoming a pressing issue due to sea level rise. Here, we assessed how a freshwater coastal wetland ecosystem responds to saltwater intrusion. In wetland mesocosms, we continuously exposed Cladium jamaicense Crantz (sawgrass) plants and their peat soil collected from a freshwater marsh to two factors associated with saltwater intrusion in karstic ecosystems: elevated loading of salinity and phosphorus (P) inputs. We took repeated measures using a 2 × 2 factorial experimental design (n = 6) with treatments composed of elevated salinity (~9 ppt), P loading (14.66 µmol P/d), or a combination of both. We measured changes in water physicochemistry, ecosystem productivity, and plant biomass change over two years to assess monthly and two-year responses to saltwater intrusion. In the short-term, plants exhibited positive growth responses with simulated saltwater intrusion (salinity + P), driven by increased P availability. Despite relatively high salinity levels for a freshwater marsh (~9 ppt), gross ecosystem productivity (GEP), net ecosystem productivity (NEP), and aboveground biomass were significantly higher in the elevated salinity + P treated monoliths compared to the freshwater controls. Salinity stress became evident after extended exposure. Although still higher than freshwater controls, GEP and NEP were significantly lower in the elevated salinity + P treatment than the +P treatment after two years. However, elevated salinity decreased live root biomass regardless of whether P was added. Our results suggest that saltwater intrusion into karstic freshwater wetlands may initially act as a subsidy by stimulating aboveground primary productivity of marsh plants. However, chronic exposure to elevated salinity results in plant stress, negatively impacting belowground peat soil structure and stability through a reduction in plant roots.

Author Correction: Warming shortens flowering seasons of tundra plant communities.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: "This work was supported by the Norwegian Research Council SnoEco project, grant number 230970". This text has now been added.

Warming shortens flowering seasons of tundra plant communities.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Author Correction: Accuracy and Precision of Tidal Wetland Soil Carbon Mapping in the Conterminous United States.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has not been fixed in the paper.

Salinity pulses interact with seasonal dry-down to increase ecosystem carbon loss in marshes of the Florida Everglades.

Coastal wetlands are globally important sinks of organic carbon (C). However, to what extent wetland C cycling will be affected by accelerated sea-level rise (SLR) and saltwater intrusion is unknown, especially in coastal peat marshes where water flow is highly managed. Our objective was to determine how the ecosystem C balance in coastal peat marshes is influenced by elevated salinity. For two years, we made monthly in situ manipulations of elevated salinity in freshwater (FW) and brackish water (BW) sites within Everglades National Park, Florida, USA. Salinity pulses interacted with marsh-specific variability in seasonal hydroperiods whereby effects of elevated pulsed salinity on gross ecosystem productivity (GEP), ecosystem respiration (ER), and net ecosystem productivity (NEP) were dependent on marsh inundation level. We found little effect of elevated salinity on C cycling when both marsh sites were inundated, but when water levels receded below the soil surface, the BW marsh shifted from a C sink to a C source. During these exposed periods, we observed an approximately threefold increase in CO2 efflux from the marsh as a result of elevated salinity. Initially, elevated salinity pulses did not affect Cladium jamaicense biomass, but aboveground biomass began to be significantly decreased in the saltwater amended plots after two years of exposure at the BW site. We found a 65% (FW) and 72% (BW) reduction in live root biomass in the soil after two years of exposure to elevated salinity pulses. Regardless of salinity treatment, the FW site was C neutral while the BW site was a strong C source (-334 to -454 g C·m-2 ·yr-1 ), particularly during dry-down events. A loss of live roots coupled with annual net CO2 losses as marshes transition from FW to BW likely contributes to the collapse of peat soils observed in the coastal Everglades. As SLR increases the rate of saltwater intrusion into coastal wetlands globally, understanding how water management influences C gains and losses from these systems is crucial. Under current Everglades' water management, drought lengthens marsh dry-down periods, which, coupled with saltwater intrusion, accelerates CO2 loss from the marsh.

Accuracy and Precision of Tidal Wetland Soil Carbon Mapping in the Conterminous United States.

Tidal wetlands produce long-term soil organic carbon (C) stocks. Thus for carbon accounting purposes, we need accurate and precise information on the magnitude and spatial distribution of those stocks. We assembled and analyzed an unprecedented soil core dataset, and tested three strategies for mapping carbon stocks: applying the average value from the synthesis to mapped tidal wetlands, applying models fit using empirical data and applied using soil, vegetation and salinity maps, and relying on independently generated soil carbon maps. Soil carbon stocks were far lower on average and varied less spatially and with depth than stocks calculated from available soils maps. Further, variation in carbon density was not well-predicted based on climate, salinity, vegetation, or soil classes. Instead, the assembled dataset showed that carbon density across the conterminous united states (CONUS) was normally distributed, with a predictable range of observations. We identified the simplest strategy, applying mean carbon density (27.0 kg C m-3), as the best performing strategy, and conservatively estimated that the top meter of CONUS tidal wetland soil contains 0.72 petagrams C. This strategy could provide standardization in CONUS tidal carbon accounting until such a time as modeling and mapping advancements can quantitatively improve accuracy and precision.

Effects of sea-level rise and freshwater management on long-term water levels and water quality in the Florida Coastal Everglades.

Since the 1880s, hydrological modification of the Greater Florida Everglades has reduced water levels and flows in Everglades National Park (ENP). The Comprehensive Everglades Restoration Program (CERP) began in 2000 to restore pre-drainage flows and preserve the natural landscape of the Everglades. However, sea-level rise (SLR) was not considered in the development of CERP. We used long-term data (2001-2016) from the Florida Coastal Everglades-Long Term Ecological Research Program to quantify and model the spatial dynamics of water levels, salinity, and nutrients in response to changes in climate, freshwater management and SLR in the Shark River Slough (SRS), ENP. Results indicate that fresh-to-marine head difference (FMHD) was the single most important factor affecting marine-to-freshwater hydrologic connectivity and transport of salinity and phosphorous upstream from the Gulf of Mexico. Sea-level has increasingly exceeded ground surface elevation at the most downstream freshwater site in SRS, thereby reducing the FMHD. We showed a higher impact of SLR in the dry season when there was practically no freshwater inflow to raise FMHD. We also demonstrated effectiveness of inflow depends more on the monthly distribution than the total annual volume. Hence, the impact per unit volume of inflow is significantly higher in the dry season in preventing high salinity and marine-derived nutrient levels. We advocate that FMHD needs to be factored into water management decisions to reduce adverse and likely irreversible effects of SLR throughout the Everglades landscape.

Coastal wetland management as a contribution to the US National Greenhouse Gas Inventory.

The IPCC 2013 Wetlands Supplement provided new guidance for countries on inclusion of wetlands in their National GHG Inventories. The United States has responded by including managed coastal wetlands for the first time in its 2017 GHG Inventory report along with an updated time series in the most recent 2018 submission and plans to update the time series on an annual basis as part of its yearly submission to the United Nations Framework Convention on Climate Change (UNFCCC). The United States followed IPCC Good Practice Guidance when reporting sources and sinks associated with managed coastal wetlands. Here we show that intact vegetated coastal wetlands are a net sink for GHGs. Despite robust regulation that has protected substantial stocks of carbon, the United States continues to lose coastal wetlands to development and the largest loss of wetlands to open water occurs around the Mississippi Delta due mostly to upstream changes in hydrology and sediment delivery, and oil and gas extraction. These processes create GHG emissions. By applying comprehensive Inventory reporting, scientists in the United States have identified opportunities for reducing GHG emissions through restoration of coastal wetlands that also provide many important societal co-benefits.

Greater temperature sensitivity of plant phenology at colder sites: implications for convergence across northern latitudes.

Warmer temperatures are accelerating the phenology of organisms around the world. Temperature sensitivity of phenology might be greater in colder, higher latitude sites than in warmer regions, in part because small changes in temperature constitute greater relative changes in thermal balance at colder sites. To test this hypothesis, we examined up to 20 years of phenology data for 47 tundra plant species at 18 high-latitude sites along a climatic gradient. Across all species, the timing of leaf emergence and flowering was more sensitive to a given increase in summer temperature at colder than warmer high-latitude locations. A similar pattern was seen over time for the flowering phenology of a widespread species, Cassiope tetragona. These are among the first results highlighting differential phenological responses of plants across a climatic gradient and suggest the possibility of convergence in flowering times and therefore an increase in gene flow across latitudes as the climate warms.

Contributions of organic and inorganic matter to sediment volume and accretion in tidal wetlands at steady state.

A mixing model derived from first principles describes the bulk density (BD) of intertidal wetland sediments as a function of loss on ignition (LOI). The model assumes that the bulk volume of sediment equates to the sum of self-packing volumes of organic and mineral components or BD = 1/[LOI/k1 + (1-LOI)/k2], where k1 and k2 are the self-packing densities of the pure organic and inorganic components, respectively. The model explained 78% of the variability in total BD when fitted to 5075 measurements drawn from 33 wetlands distributed around the conterminous United States. The values of k1 and k2 were estimated to be 0.085 ± 0.0007 g cm-3 and 1.99 ± 0.028 g cm-3, respectively. Based on the fitted organic density (k1) and constrained by primary production, the model suggests that the maximum steady state accretion arising from the sequestration of refractory organic matter is ≤ 0.3 cm yr-1. Thus, tidal peatlands are unlikely to indefinitely survive a higher rate of sea-level rise in the absence of a significant source of mineral sediment. Application of k2 to a mineral sediment load typical of East and eastern Gulf Coast estuaries gives a vertical accretion rate from inorganic sediment of 0.2 cm yr-1. Total steady state accretion is the sum of the parts and therefore should not be greater than 0.5 cm yr-1 under the assumptions of the model. Accretion rates could deviate from this value depending on variation in plant productivity, root:shoot ratio, suspended sediment concentration, sediment-capture efficiency, and episodic events.

Global assessment of experimental climate warming on tundra vegetation: heterogeneity over space and time.

Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Fire and grazing in a mesic tallgrass prairie: impacts on plant species and functional traits.

Fire is a globally distributed disturbance that impacts terrestrial ecosystems and has been proposed to be a global "herbivore." Fire, like herbivory, is a top-down driver that converts organic materials into inorganic products, alters community structure, and acts as an evolutionary agent. Though grazing and fire may have some comparable effects in grasslands, they do not have similar impacts on species composition and community structure. However, the concept of fire as a global herbivore implies that fire and herbivory may have similar effects on plant functional traits. Using 22 years of data from a mesic, native tallgrass prairie with a long evolutionary history of fire and grazing, we tested if trait composition between grazed and burned grassland communities would converge, and if the degree of convergence depended on fire frequency. Additionally, we tested if eliminating fire from frequently burned grasslands would result in a state similar to unburned grasslands, and if adding fire into a previously unburned grassland would cause composition to become more similar to that of frequently burned grasslands. We found that grazing and burning once every four years showed the most convergence in traits, suggesting that these communities operate under similar deterministic assembly rules and that fire and herbivory are similar disturbances to grasslands at the trait-group level of organization. Three years after reversal of the fire treatment we found that fire reversal had different effects depending on treatment. The formerly unburned community that was then burned annually became more similar to the annually burned community in trait composition suggesting that function may be rapidly restored if fire is reintroduced. Conversely, after fire was removed from the annually burned community trait composition developed along a unique trajectory indicating hysteresis, or a time lag for structure and function to return following a change in this disturbance regime. We conclude that functional traits and species-based metrics should be considered when determining and evaluating goals for fire management in mesic grassland ecosystems.

Modeling Nymphoides architecture: A morphological analysis of Nymphoides aquatica (Menyanthaceae).

PREMISE OF THE STUDY: Species in the aquatic genus Nymphoides have inflorescences that appear to arise from the petioles of floating leaves. The inflorescence-floating leaf complex can produce vegetative propagules and/or additional inflorescences and leaves. We analyzed the morphology of N. aquatica to determine how this complex relates to whole plant architecture and whether whole plant growth is sympodial or monopodial. • METHODS: We used dissections, measurements, and microscopic observations of field-collected plants and plants cultivated for 2 years in outdoor tanks in south Florida, USA. • KEY RESULTS: Nymphoides aquatica had a submerged plagiotropic rhizome that produced floating leaves in an alternate/spiral phyllotaxy. Rhizomes were composed of successive sympodial units that varied in the number of leaves produced before the apex terminated. The basic sympodial unit had a prophyll that subtended a renewal-shoot bud, a short-petioled leaf (SPL) with floating lamina, and an inflorescence; the SPL axillary bud expanded as a vegetative propagule. Plants produced either successive basic sympodial units or expanded sympodia that intercalated long-petioled leaves between the prophyll and the SPL. • CONCLUSIONS: Nymphoides aquatica grows sympodially, forming a rhizome composed of successive basic sympodia and expanded sympodial units. Variations on these types of sympodial growth help explain the branching patterns and leaf morphologies described for other Nymphoides species. Monitoring how these two sympodial phases are affected by water depth provides an ecologically meaningful way to assess N. aquatica's responses to altered hydrology.