Comparative skeletal anatomy of salt marsh and western harvest mice in relation to locomotor ecology.

The salt marsh harvest mouse (Reithrodontomys raviventris) is an endangered species, endemic to the San Francisco Bay Estuary, that co-occurs with the more broadly distributed species, the western harvest mouse (Reithrodontomys megalotis). Despite their considerable external morphological similarities, the northern subspecies of salt marsh harvest mice have relatively longer and thicker tails than do western harvest mice, which may be related to their abilities to climb emergent marsh vegetation to avoid tidal inundation. We used micro-CT to compare post-cranial skeletal anatomy between the salt marsh and western harvest mouse, to examine whether the salt marsh harvest mouse's restriction to brackish marshes is associated with skeletal adaptations for scansorial locomotion. We found that salt marsh harvest mice exhibited a deeper 3rd caudal vertebra, a more caudally located longest tail vertebra, craniocaudally longer tail vertebrae, and a longer digit III proximal phalanx than western harvest mice. These phalangeal and vertebral characteristics are known to decrease body rotations during climbing, increase contact with substrates, and decrease fall susceptibility in arboreal mammals, suggesting that the salt marsh harvest mouse may be morphologically specialized for scansorial locomotion, adaptive for its dynamic wetland environment.

High methane concentrations in tidal salt marsh soils: Where does the methane go?

Tidal salt marshes produce and emit CH4 . Therefore, it is critical to understand the biogeochemical controls that regulate CH4 spatial and temporal dynamics in wetlands. The prevailing paradigm assumes that acetoclastic methanogenesis is the dominant pathway for CH4 production, and higher salinity concentrations inhibit CH4 production in salt marshes. Recent evidence shows that CH4 is produced within salt marshes via methylotrophic methanogenesis, a process not inhibited by sulfate reduction. To further explore this conundrum, we performed measurements of soil-atmosphere CH4 and CO2 fluxes coupled with depth profiles of soil CH4 and CO2 pore water gas concentrations, stable and radioisotopes, pore water chemistry, and microbial community composition to assess CH4 production and fate within a temperate tidal salt marsh. We found unexpectedly high CH4 concentrations up to 145,000 µmol mol-1 positively correlated with S2- (salinity range: 6.6-14.5 ppt). Despite large CH4 production within the soil, soil-atmosphere CH4 fluxes were low but with higher emissions and extreme variability during plant senescence (84.3 ± 684.4 nmol m-2 s-1 ). CH4 and CO2 within the soil pore water were produced from young carbon, with most Δ14 C-CH4 and Δ14 C-CO2 values at or above modern. We found evidence that CH4 within soils was produced by methylotrophic and hydrogenotrophic methanogenesis. Several pathways exist after CH4 is produced, including diffusion into the atmosphere, CH4 oxidation, and lateral export to adjacent tidal creeks; the latter being the most likely dominant flux. Our findings demonstrate that CH4 production and fluxes are biogeochemically heterogeneous, with multiple processes and pathways that can co-occur and vary in importance over the year. This study highlights the potential for high CH4 production, the need to understand the underlying biogeochemical controls, and the challenges of evaluating CH4 budgets and blue carbon in salt marshes.

Las marismas salinas producen y emiten CH4 . Por lo tanto, es esencial comprender los controles biogeoquímicos que regulan la dinámica espacial y temporal del CH4 en estos humedales. El paradigma predominante asume que la metanogénesis acetoclástica es la vía dominante para la producción de CH4 y que altas concentraciones de salinidad inhiben la producción de CH4 en estos ecosistemas. Hay evidencia que el CH4 se produce las marismas salinas a través de la metanogénesis metilotrófica, un proceso no inhibido por la reducción del sulfato. Para explorar esta paradoja, realizamos mediciones de los flujos de CH4 y CO2 del suelo a la atmósfera junto con perfiles de concentraciones de CH4 y CO2 en el suelo, isótopos estables y radioisótopos, química del agua y composición de la comunidad microbiana para evaluar la producción y el destino del CH4 en una marisma salina templada. Encontramos concentraciones de CH4 sorprendentemente altas de hasta 145,000 µmol mol−1 correlacionadas positivamente con S2− (rango de salinidad: 6.6 a 14.5 ppt). A pesar de la gran producción de CH4 en el suelo, los flujos de CH4 del suelo a la atmósfera fueron bajos, pero con mayores emisiones y variabilidad extrema durante la época de senescencia de las plantas (84.3 ± 684.4 nmol m−2 s−1 ). El CH4 y el CO2 en el suelo se produjeron a partir de carbono joven, con la mayoría de los valores Δ14 C-CH4 y Δ14 C-CO2 en o por encima de valores modernos. Encontramos evidencia de que el CH4 en los suelos fue producido por metanogénesis metilotrófica e hidrogenotrófica. Existen varias vías que el CH4 producido sigue, incluida la difusión hacia la atmósfera, la oxidación del CH4 y la exportación lateral a arroyos adyacentes a la marisma; siendo este último el flujo dominante más probable. Nuestros hallazgos demuestran que la producción y los flujos de CH4 son biogeoquímicamente heterogéneos, con múltiples procesos y vías que pueden coexistir y variar en importancia a lo largo del año. Este estudio destaca el potencial de alta producción de CH4 , la necesidad de comprender los controles biogeoquímicos de la producción de CH4 y los retos que existen para evaluar las reservas de CH4 y el carbono azul en marismas salinas.

Tidal channel meanders serve as stepping-stones to facilitate cordgrass landward spread by creating invasion windows.

Understanding the mechanisms by which the geomorphic structures affect habitat invasibility by mediating various abiotic and biotic factors is essential for predicting whether these geomorphic structures may provide spatial windows of opportunity to facilitate range-expansion of invasive species in salt marshes. Many studies have linked geomorphic landscape features such as tidal channels to invasion by exotic plants, but the role of tidal channel meanders (i.e., convex and concave sides) in regulating the Spartina alterniflora invasion remains unclear. Here, we examined the combined effects of tidal channel meander-mediated hydrodynamic variables, soil abiotic stresses, and propagule pressure on the colonization of Spartina in the Yellow River Delta, China, by conducting field observations and experiments. The results showed that lower hydrodynamic disturbance, bed shear stress, and higher propagule pressure triggered by eddies due to the convex structure of channel meanders facilitated Spartina seedling establishment and growth, whereas the concave side considerably inhibited the Spartina invasion. Lower soil abiotic stresses also significantly promoted the invasibility of the channel meanders by Spartina. Based on these findings, we propose a conceptual framework to illustrate the effects of the meandering geomorphology of tidal channels on the mechanisms that might allow the landward spread of Spartina and related processes. Our results demonstrate that the meandering geomorphic structures of tidal channels could act as stepping-stones to significantly facilitate the landward invasion of Spartina along tidal channels. This implies that geomorphic characteristics of tidal channels should be integrated into invasive species control and salt marsh management strategies.

Role of n-DAMO in Mitigating Methane Emissions from Intertidal Wetlands Is Regulated by Saltmarsh Vegetations.

Coastal wetlands are hotspots for methane (CH4) production, reducing their potential for global warming mitigation. Nitrite/nitrate-dependent anaerobic methane oxidation (n-DAMO) plays a crucial role in bridging carbon and nitrogen cycles, contributing significantly to CH4 consumption. However, the role of n-DAMO in reducing CH4 emissions in coastal wetlands is poorly understood. Here, the ecological functions of the n-DAMO process in different saltmarsh vegetation habitats as well as bare mudflats were quantified, and the underlying microbial mechanisms were explored. Results showed that n-DAMO rates were significantly higher in vegetated habitats (Scirpus mariqueter and Spartina alterniflora) than those in bare mudflats (P < 0.05), leading to an enhanced contribution to CH4 consumption. Compared with other habitats, the contribution of n-DAMO to the total anaerobic CH4 oxidation was significantly lower in the Phragmites australis wetland (15.0%), where the anaerobic CH4 oxidation was primarily driven by ferric iron (Fe3+). Genetic and statistical analyses suggested that the different roles of n-DAMO in various saltmarsh wetlands may be related to divergent n-DAMO microbial communities as well as environmental parameters such as sediment pH and total organic carbon. This study provides an important scientific basis for a more accurate estimation of the role of coastal wetlands in mitigating climate change.

Colonization of mudflat substrate by microarthropods: the role of distance, inundation frequency and body size.

Salt marshes represent a unique ecosystem at the marine-terrestrial boundary of shallow protected coastlines. Microarthropods form an essential component of soil food webs, but how they colonize new intertidal habitats is little understood. By establishing two experimental systems without animals, we investigated microarthropod colonization (1) at the seashore from the pioneer zone to the lower and upper salt marsh and (2) at the same tidal height on artificial islands 500 m from the seashore. Potential source populations of microarthropods in the respective zones were also investigated. Colonization of microarthropods after 5 years was consistently faster on the seashore than on the artificial islands. Collembola and Mesostigmata colonized all the zones both on the seashore and on the artificial islands, with colonization being faster in the upper salt marsh and in the pioneer zone than in the lower salt marsh. Oribatida colonized the new habitats on the seashore, but only little on the artificial islands. Variations in species composition were more pronounced between salt marsh zones than between experimental systems, indicating that local environmental conditions (i.e., inundation frequency) are more important for the assembly of microarthropod communities than the distance from source populations (i.e., dispersal processes). Variations in community body size of Oribatida and Mesostigmata indicated environmental filtering of traits, with smaller species suffering from frequent inundations. Notably, Mesostigmata most successfully colonized the new habitats across salt marsh zones on both systems. Overall, the results document major mechanisms of colonization of intertidal habitats by microarthropods with different life histories and feeding strategies.

Global potential distribution of mangroves: Taking into account salt marsh interactions along latitudinal gradients.

Mangrove is one of the most productive and sensitive ecosystems in the world. Due to the complexity and specificity of mangrove habitat, the development of mangrove is regulated by several factors. Species distribution models (SDMs) are effective tools to identify the potential habitats for establishing and regenerating the ecosystem. Such models usually include exclusively environmental factors. Nevertheless, recent studies have challenged this notion and highlight the importance of including biotic interactions. Both factors are necessary for a mechanistic understanding of the mangrove distribution in order to promote the protection and restoration of mangroves. Thus, we present a novel approach of combining environmental factors and interactions with salt marsh for projecting mangrove distributions at the global level and within latitudinal zones. To test the salt marsh interaction, we fit the MaxEnt model with two predicting sets: (1) environments only and (2) environments + salt marsh interaction index (SII). We found that both sets of models had good predictive ability, although the SII improved model performance slightly. Potential distribution areas of mangrove decrease with latitudes, and are controlled by biotic and abiotic factors. Temperature, precipitation and wind speed are generally critical at both global scale and ecotones along latitudes. SII is important on global scale, with a contribution of 5.9%, ranking 6th, and is particularly critical in the 10-30°S and 20-30°N zone. Interactions with salt marsh, including facilitation and competition, are shown to affect the distribution of mangroves at the zone of coastal ecotone, especially in the latitudinal range from 10° - 30°. The contribution of SII to mangrove distribution increases with latitudes due to the difference in the adaptive capacity of salt marsh plants and mangroves to environments. Totally, this study identified and quantified the effects of salt marsh on mangrove distribution by establishing the SII. The results not only facilitate to establish a more accurate mangrove distribution map, but also improve the efficiency of mangrove restoration by considering the salt marsh interaction in the mangrove management projects. In addition, the method of incorporating biotic interaction into SDMs through establish the biotic interaction index has contributed to the development of SDMs.

Precipitation changes alter plant dominant species and functional groups by changing soil salinity in a coastal salt marsh.

Specific mechanisms of precipitation change due to global climate variability on plant communities in coastal salt marsh ecosystems remain unknown. Hence, a field manipulative precipitation experiment was established in 2014 and 5 years of field surveys of vegetation from 2017 to 2021 to explore the effects of precipitation changes on plant community composition. The results showed that changes in plant community composition were driven by dominant species, and that the dominance of key species changed significantly with precipitation gradient and time, and that these changes ultimately altered plant community traits (i.e., community density, height, and species richness). Community height increased but community density decreased with more precipitation averaged five years. Furthermore, changes in precipitation altered dominant species composition and functional groups mainly by influencing soil salinity. Salinity stress caused by decreased precipitation shifted species composition from a dominance of taller perennials and grasses to dwarf annuals and forbs, while the species richness decreased. Conversely, soil desalination caused by increased precipitation increased species richness, especially increasing in the dominance of grasses and perennials. Specifically, Apocynaceae became dominance from rare while Amaranthaceae decreased in response to increased precipitation, but Poaceae was always in a position of dominance. Meanwhile, the dominance of grasses and perennials has the cumulative effect of years and their proportion increased under the increased 60% of ambient precipitation throughout the years. However, the annual forb Suaeda glauca was gradually losing its dominance or even becoming extinct over years. Our study highlights that the differences in plant salinity tolerance are key to the effects of precipitation changes on plant communities in coastal salt marsh. These findings aim to provide a theoretical basis for predicting vegetation dynamics and developing ecological management strategies to adapt to future precipitation changes.

Succession of Fungal Communities and Their Functional Profiles in a Decaying Foundation Species.

Dead plant biomass from foundation plant species is fundamental for the survival of coastal salt marshes because dead biomass aids in the vertical accretion of the ecosystem. Fungi regulate the decomposition of dead biomass, and thus play an essential role for marsh survival. Despite their importance, little is known about the compositional and functional changes of fungal communities in plant matter throughout senescence and litter decomposition. Here, we analyzed how fungal communities and their functionality change in the foundation plant species Spartina patens, which builds vast amounts of dead plant biomass (thatch) on the soil surface. We analyzed the chemical and fungal properties of live biomass, standing dead biomass (dead biomass shortly after senescence), upper thatch (top layer of litter on the soil surface), and lower thatch (bottom layer of litter on the soil surface) during September and November of 2021. We found that the chemical and fungal properties of different S. patens biomass types followed a predictable litter decomposition pattern. Notably, live biomass, standing dead biomass, upper thatch, and lower thatch all hosted unique fungal communities and litter chemistry. Functional groups present in live biomass (pathogens, epiphytes, and mycoparasites) were lost during senescence and later replaced by diverse saprotrophs. The abundance of lignocellulose saprotrophs increased throughout decomposition, with the highest abundance occurring in lower thatch. These results suggest a predictable succession of fungal communities through the senescence and decomposition of the foundation species S. patens. Our study highlights the diversity of fungal communities in a disappearing foundation species.

Microbial Community Succession Along a Chronosequence in Constructed Salt Marsh Soils.

In this study, we examined the succession of soil microbial communities across a chronosequence of newly constructed salt marshes constructed primarily of fine-grained dredge material, using 16S rRNA amplicon sequences. Alpha diversity in the subsurface horizons was initially low and increased to reference levels within 3 years of marsh construction, while alpha diversity in the newly accumulating organic matter-rich surface soils was initially high and remained unchanged. Microbial community succession was fastest in the surface horizon (~ 24 years to reference equivalency) and became progressively slower with depth in the subsurface horizons (~ 30-67 years). Random forest linear regression analysis was used to identify important taxa driving the trajectories toward reference conditions. In the parent material, putative sulfate-reducers (Desulfobacterota), methanogens (Crenarchaeota, especially Methanosaeta), and fermenters (Chloroflexi and Clostridia) increased over time, suggesting an enrichment of these metabolisms over time, similar to natural marshes. Concurrently in the surface soils, the relative abundances of putative methane-, methyl-, and sulfide oxidizers, especially among Gammaproteobacteria, increased over time, suggesting the co-development of sulfide and methane removal metabolisms in marsh soils. Finally, we observed that the surface soil communities at one of the marshes did not follow the trajectory of the others, exhibiting a greater relative abundance of anaerobic taxa. Uniquely in this dataset, this marsh was developing signs of excessive inundation stress in terms of vegetation coverage and soil geochemistry. Therefore, we suggest that soil microbial community structure may be effective bioindicators of salt marsh inundation and are worthy of further targeted investigation.

Non-native Plant Species Invasion Increases the Importance of Deterministic Processes in Fungal Community Assembly in a Coastal Wetland.

Fungal communities are essential to the maintenance of soil multifunctionality. Plant invasion represents a growing challenge for the conservation of soil biodiversity across the globe, but the impact of non-native species invasion on fungal diversity, community structure, and assembly processes remains largely unknown. Here, we examined the diversity, community composition, functional guilds, and assembly process of fungi at three soil depths underneath a native species, three non-native species, and a bare tidal flat from a coastal wetland. Plant species was more important than soil depth in regulating the diversity, community structure, and functional groups of fungi. Non-native species, especially Spartina alterniflora, increased fungal diversity, altered fungal community structure, and increased the relative abundance of saprotrophic and pathogenic fungi in coastal wetland soils. Stochastic processes played a predominant role in driving fungal community assembly, explaining more than 70% of the relative contributions. However, compared to a native species, non-native species, especially S. alterniflora, reduced the relative influence of stochastic processes in fungal community assembly. Collectively, our results provide novel evidence that non-native species can increase fungal diversity, the relative abundance of saprotrophic and pathogenic fungi, and deterministic processes in the assembly of fungi in coastal wetlands, which can expand our knowledge of the dynamics of fungal communities in subtropical coastal wetlands.

Hazardous and contaminated sites within salt marsh migration corridors in Rhode Island, USA.

As salt marshes attempt to migrate upland due to sea level rise, they will encounter many kinds of land development and infrastructure in highly populated, urbanized coastal communities. Hazardous and contaminated sites (HCSs) -- facilities and infrastructure that store, use, or release harmful substances -- are particularly concerning obstacles to salt marsh migration because of their potential to release contaminants if their structural integrity is compromised. Inventorying HCSs within migration pathways can inform coastal resilience planning. To understand what kinds of HCSs migrating marsh may encounter in Rhode Island, USA, we inventoried sites from federal and state sources, assigned contaminant hazard rankings to most sites, and overlayed them with projected marsh migration corridors. We found that HCSs are extensive across marsh migration corridors in the state, especially in urban areas. Among the most common HCSs in and around Rhode Island salt marshes are stormwater outfalls, underground storage tanks, and facilities registered with EPA's Resource Conservation and Recovery Act (RCRA) or EPA's National Pollutant Discharge Elimination System (NPDES). These sites pose varying hazards to human and aquatic life if breached, with some sites representing little or no threat but most posing some degree of hazard to their surroundings. This coastal HCSs inventory can inform prioritization and management of coastal salt marshes subject to accelerated sea level rise. Management decisions such as allowing marsh migration, implementing adaptation actions to build salt marsh elevation, or erecting physical barriers at marsh sites will influence future salt marsh extent, marshes' ability to provide ecosystem services, and public health exposures to toxic releases. In addition, as Rhode Island and other coastal states work to promote coastal resiliency, this type of inventory can inform decisions about which HCSs to prioritize for remediation and other climate adaptation actions. Marsh migration is just one potential consequence of sea level rise, so many of the considerations outlined here are widely applicable to the broader goal of preparing coastal communities for rising seas.

The importance of structural and functional characteristics of tidal channels to smooth cordgrass invasion in the Yellow River Delta, China: Implications for coastal wetland management.

Understanding the spatiotemporal landscape dynamics and spread pathways of invasive plants, as well as their interactions with geomorphic landscape features, are of great importance for predicting and managing their future range-expansion in non-native habitats. Although previous studies have linked geomorphic landscape features such as tidal channels to plant invasions, the potential mechanisms and critical characteristics of tidal channels that affect the landward invasion by Spartina alterniflora, an aggressive plant in global coastal wetlands, remain unclear. Here, using high-resolution remote-sensing images of the Yellow River Delta from 2013 to 2020, we first quantified the evolution of tidal channel networks by analyzing the spatiotemporal dynamics of their structural and functional characteristics. The invasion patterns and pathways of S. alterniflora were then identified. Based on the above-mentioned quantification and identification, we finally quantified the influences of tidal channel characteristics on S. alterniflora invasion. The results showed that tidal channel networks presented increasing growth and development over time, and their spatial structure evolved from simple to complex. The external isolated expansion of S. alterniflora played a dominant role during the initial invasion stage, and then they connected the discrete patches into the meadow through marginal expansion. Afterwards, tidal channel-driven expansion gradually increased and became the primary way during the late invasion stage, accounting for about 47.3%. Notably, tidal channel networks with higher drainage efficiency (shorter OPL, higher D and E) attained larger invasion areas. The longer the tidal channels and the more sinuous the channel structure, the greater the invasion potential by S. alterniflora. These findings highlight the importance of structural and functional properties of tidal channel networks in driving plant invasion landward, which should be incorporated into future control and management of invasive plants in coastal wetlands.

Assessing long-term outcomes of tidal restoration in New England salt marshes.

Salt marshes are valuable coastal ecosystems, but many have been degraded by roads, railways, and other infrastructure that restrict tidal flow and impound watershed runoff. Restoration of tidal flow to tide-restricted salt marshes generally aims to restore native vegetation and habitat functions. Biological communities may take one or more decades to recover following tidal restoration, but outcomes are seldom assessed on that timescale. We assessed the long-term outcomes of eight tidal restorations in Rhode Island, USA using observed changes in plant and nekton communities from pre-restoration to present, and newly-collected data from a rapid assessment method. The time-series vegetation and nekton data suggest that while restoration actions promoted biological recovery, ambient factors such as inundation stress and eutrophication have worked to offset it. Rapid assessment results indicate that the cover of Phragmites australis is higher and the cover of meadow high marsh is lower at restoration marshes compared with a broad reference sample, suggesting incomplete recovery on average, although outcomes varied across the restoration marshes. Habitat integrity increased with the degree of adaptive management following restoration, as well as the age of restoration, but salt marsh restoration practitioners may need to shift their methods and expectations to accommodate human influences on ambient environmental conditions, particularly prevalent, increasing inundation stress associated with sea-level rise. Our study highlights the value of standardized long-term biological monitoring in assessing salt marsh restoration outcomes, and demonstrates how rapid assessment data can add valuable context to restoration findings.

Quantifying the Impact of Hydrological Connectivity on Salt Marsh Vegetation in the Liao River Delta Wetland.

Salt marshes play a critical role in ecological functioning and have significant economic value. Hydrological elements are considered to be one of the major contributors to salt marsh degradation. However, how hydrological connectivity affects salt marshes remains poorly investigated at fine scales. This paper used spatial analysis and statistical methods to investigate the impact of hydrological connectivity on the spatial and temporal distribution characteristics of salt marsh vegetation in two natural succession areas of the Liao River Delta wetland in 2020 and 2021 by selecting vegetation area, NDVI, tidal creeks area, distance to tidal creeks, and the Index of Connectivity, using 1 m Gaofen-2 data and 0.2 m aerial topographic data. The study found that the area and growth status of vegetation and the overall connectivity in 2021 were better than that in 2020, while the west bank of the Liao River was better than that on the east bank. Phragmites australis showed a round island distribution pattern primarily at the end of tidal creeks. The differences between different hydrological connectivity and vegetation area were significant in 2021. The vegetation area was the largest under poor and moderate connectivity. We also found that within a distance range of 0-6 m from tidal creeks, the vegetation area increased with increasing distance, but beyond 6 m, the vegetation area decreased with increasing distance. Our results showed that poor and moderate connectivity conditions were more suitable for vegetation growth. The threshold value of 6 m can provide an important reference for wetland vegetation restoration in the Liao River Delta wetland. Supplementary Information: The online version contains supplementary material available at 10.1007/s13157-023-01693-4.

Exploring the Use of Living Shorelines for Stabilization and Nutrient Mitigation in New England.

As salt marsh habitats face challenges due to sea level rise, storm events, and coastal development, there is an effort to use nature-based approaches such as living shorelines to enhance salt marshes and provide coastal protection. A living shoreline restoration and seasonal monitoring was conducted between July 2016 and October 2018 at an eroding salt marsh on Martha's Vineyard, Massachusetts, Northeastern USA to assess changes in two essential ecosystem services: shoreline stabilization and nitrogen removal. Neither the living shoreline nor unaltered sites demonstrated significant sediment deposition at the marsh edge or on the marsh platform between 2017 and 2018. While we expected nitrogen removal via denitrification to improve at the living shoreline sites over time as abiotic and biotic conditions became more favorable, we found limited support for this hypothesis. We found higher rates of denitrification enzyme activity (DEA) at the living shoreline sites when compared to unaltered sites, but these rates did not increase over time. This study also provides a qualitative assessment of our living shoreline structural integrity through the years, particularly following storm events that greatly challenged our restoration efforts. We demonstrate that living shorelines fortified solely with natural materials may not be the most effective approach to maintain these ecosystem services for Northeastern USA salt marshes exposed to intense northeasterly storms. We suggest the restoration of salt marshes to improve major functions be a priority among managers and restoration practitioners. Initiatives promoting the use of nature-based restoration solution where environmental conditions permit should be encouraged.

Biota-mediated carbon cycling-A synthesis of biotic-interaction controls on blue carbon.

Research into biotic interactions has been a core theme of ecology for over a century. However, despite the obvious role that biota play in the global carbon cycle, the effects of biotic interactions on carbon pools and fluxes are poorly understood. Here we develop a conceptual framework that illustrates the importance of biotic interactions in regulating carbon cycling based on a literature review and a quantitative synthesis by means of meta-analysis. Our study focuses on blue carbon ecosystems-vegetated coastal ecosystems that function as the most effective long-term CO2 sinks of the biosphere. We demonstrate that a multitude of mutualistic, competitive and consumer-resource interactions between plants, animals and microbiota exert strong effects on carbon cycling across various spatial scales ranging from the rhizosphere to the landscape scale. Climate change-sensitive abiotic factors modulate the strength of biotic-interaction effects on carbon fluxes, suggesting that the importance of biota-mediated carbon cycling will change under future climatic conditions. Strong effects of biotic interactions on carbon cycling imply that biosphere-climate feedbacks may not be sufficiently represented in current Earth system models. Inclusion of new functional groups in these models, and new approaches to simplify species interactions, may thus improve the predictions of biotic effects on the global climate.

Plant invasion reconstructs soil microbial assembly and functionality in coastal salt marshes.

Microbiologically driven ecosystem processes can be profoundly altered by alien plant invasions. There is limited understanding of the ecological mechanisms orchestrating different microbial constituents and their roles in emerging functional properties under plant invasions. Here, we investigated soil microbial communities and functions using high-throughput amplicon sequencing and GeoChip technology, respectively, along a chronological gradient of smooth cordgrass invasion in salt marshes located in the Yellow River Estuary, China. We found a positive correlation between microbial diversity and the duration age of invasion, and both bacterial and fungal communities showed consistent changes with invasion. Soil microbial metabolic potential, as indicated by the abundance of microbial functional genes involved in biogeochemical cycling, decreased in response to invasion. As a consequence, declining soil microbial metabolisms as a result of plant invasion facilitated carbon accumulation in invaded salt marshes. Bacteria and fungi exhibited distinct contributions to assembly processes along the invasion gradient: bacterial communities were mainly driven by selection and dispersal limitation, while fungi were dramatically shaped by stochastic processes. Soil microbial-mediated functions were taxon-specific, as indicated by community-function relationships. This study demonstrates the distinct contributions of microbial constituents to microbial community assembly and functions and sheds light on the implications of plant invasion on microbiologically driven ecosystem processes in coastal wetlands.

Storage, patterns and influencing factors for soil organic carbon in coastal wetlands of China.

Soil organic carbon (SOC) in coastal wetlands, also known as "blue C," is an essential component of the global C cycles. To gain a detailed insight into blue C storage and controlling factors, we studied 142 sites across ca. 5000 km of coastal wetlands, covering temperate, subtropical, and tropical climates in China. The wetlands represented six vegetation types (Phragmites australis, mixed of P. australis and Suaeda, single Suaeda, Spartina alterniflora, mangrove [Kandelia obovata and Avicennia marina], tidal flat) and three vegetation types invaded by S. alterniflora (P. australis, K. obovata, A. marina). Our results revealed large spatial heterogeneity in SOC density of the top 1-m ranging 40-200 Mg C ha-1 , with higher values in mid-latitude regions (25-30° N) compared with those in both low- (20°N) and high-latitude (38-40°N) regions. Vegetation type influenced SOC density, with P. australis and S. alterniflora having the largest SOC density, followed by mangrove, mixed P. australis and Suaeda, single Suaeda and tidal flat. SOC density increased by 6.25 Mg ha-1 following S. alterniflora invasion into P. australis community but decreased by 28.56 and 8.17 Mg ha-1 following invasion into K. obovata and A. marina communities. Based on field measurements and published literature, we calculated a total inventory of 57 × 106 Mg C in the top 1-m soil across China's coastal wetlands. Edaphic variables controlled SOC content, with soil chemical properties explaining the largest variance in SOC content. Climate did not control SOC content but had a strong interactive effect with edaphic variables. Plant biomass and quality traits were a minor contributor in regulating SOC content, highlighting the importance of quantity and quality of OC inputs and the balance between production and degradation within the coastal wetlands. These findings provide new insights into blue C stabilization mechanisms and sequestration capacity in coastal wetlands.

Vegetation zones as indicators of denitrification potential in salt marshes.

Salt marsh vegetation zones shift in response to large-scale environmental changes such as sea-level rise (SLR) and restoration activities, but it is unclear if they are good indicators of soil nitrogen removal. Our goal was to characterize the relationship between denitrification potential and salt marsh vegetation zones in tidally restored and tidally unrestricted coastal marshes, and to use vegetation zones to extrapolate how SLR may influence high marsh denitrification at the landscape scale. We conducted denitrification enzyme activity assays on sediment collected from three vegetation zones expected to shift in distribution due to SLR and tidal flow restoration across 20 salt marshes in Connecticut, USA (n = 60 sampling plots) during the summer of 2017. We found lower denitrification potential in short-form Spartina alterniflora zones (mean, 95% CI: 4, 3-6 mg N h-1  m-2 ) than in S. patens (25, 15-36 mg N h-1  m-2 ) and Phragmites australis (56, 16-96 mg N h-1  m-2 ) zones. Vegetation zone was the single best predictor and explained 52% of the variation in denitrification potential; incorporating restoration status and soil characteristics (soil salinity, moisture, and ammonium) did not improve model fit. Because denitrification potential did not differ between tidally restored and unrestricted marshes, we suggest landscape-scale changes in denitrification after tidal restoration are likely to be associated with shifts in vegetation, rather than differences driven by restoration status. Sea-level-rise-induced hydrologic changes are widely observed to shift high marsh dominated by S. patens to short-form S. alterniflora. To explore the implications of this shift in dominant high marsh vegetation, we paired our measured mean denitrification potential rates with projections of high marsh loss from SLR. We found that, under low and medium SLR scenarios, predicted losses of denitrification potential due to replacement of S. patens by short-form S. alterniflora were substantially larger than losses due to reduced high marsh land area alone. Our results suggest that changes in vegetation zones can serve as landscape-scale predictors of the response of denitrification rates to rapid changes occurring in salt marshes.

Reef design and site hydrodynamics mediate oyster restoration and marsh stabilization outcomes.

The detrimental ecological impacts of engineered shoreline protection methods (e.g., seawalls) and the need to protect the coastal zone have prompted calls for greater use of natural and nature-based infrastructure (NNBI). To balance competing needs of structural stability and ecological functioning, managers require assessments of NNBI designs and materials for differing environmental settings (e.g., among wave-energy regimes). To examine the effects of setting and oyster-based NNBI design on the provision of shoreline protection, we constructed reefs from two substrates: a novel, biodegradable material (Oyster Catcher, OC) and traditional oyster shell bags (SB) on low- and high-energy eroding salt marsh shorelines, designated based on fetch and boat wake exposure. Both reef types buffered marsh elevation change on the high-energy shoreline relative to unaltered controls, but only SB reefs were able to do so on the low-energy shoreline. Additionally, both shorelines experienced high ambient rates of retreat and declines in marsh vegetation shoot density. Although constructed reefs did not mitigate marsh retreat on the low-energy shoreline, novel OC reefs significantly reduced retreat relative to SB reefs and control sites (no reefs) on the high-energy shoreline. Those SB reefs were severely damaged by storm events, increasing their areal footprints at the expense of vertical relief. Conversely, OC reefs on both shorelines exhibited steady oyster recruitment and growth and hosted higher densities of larger oysters. To successfully provide shoreline stabilization benefits, oyster-based NNBI must be structurally stable and able to promote sustained oyster recruitment and growth. Our results indicate that deliberate decisions regarding NNBI substrate, siting, and configuration can produce resilient reefs, which reduce rates of erosion and, in some cases, enhance vertical accretion along salt marsh edges. The growth trajectory, structural stability, and co-benefit provisioning of OC reefs demonstrate the potential of alternative restoration substrates to provide valuable oyster habitat along threatened marsh shorelines.