Global dataset of soil organic carbon in tidal marshes.

Tidal marshes store large amounts of organic carbon in their soils. Field data quantifying soil organic carbon (SOC) stocks provide an important resource for researchers, natural resource managers, and policy-makers working towards the protection, restoration, and valuation of these ecosystems. We collated a global dataset of tidal marsh soil organic carbon (MarSOC) from 99 studies that includes location, soil depth, site name, dry bulk density, SOC, and/or soil organic matter (SOM). The MarSOC dataset includes 17,454 data points from 2,329 unique locations, and 29 countries. We generated a general transfer function for the conversion of SOM to SOC. Using this data we estimated a median (± median absolute deviation) value of 79.2 ± 38.1 Mg SOC ha-1 in the top 30 cm and 231 ± 134 Mg SOC ha-1 in the top 1 m of tidal marsh soils globally. This data can serve as a basis for future work, and may contribute to incorporation of tidal marsh ecosystems into climate change mitigation and adaptation strategies and policies.

Increased extent of waterfowl grazing lengthens the recovery time of a colonizing seagrass (Halophila ovalis) with implications for seagrass resilience.

Herbivore distributions and abundance are shifting because of climate change, leading to intensified grazing pressure on foundation species such as seagrasses. This, combined with rapidly increasing magnitudes of change in estuarine ecosystems, may affect seagrass resilience. While the overall resilience of seagrasses is generally well-studied, the timeframes of recovery has received comparatively little attention, particularly in temperate estuaries. We investigated how the recovery time (RT) of seagrass is affected by simulated grazing in a southwestern Australian estuary. Whilst excluding swans, we simulated different grazing intensities (25, 50, 75, and 100% removal from 1 m2 plots) at four locations in the Swan-Canning Estuary, Western Australia during summer and tracked the recovery of seagrass over 3 months, using seagrass cover as the main measure of recovery. We found that seagrass recovered within 4-6 weeks from the lower grazing intensities (25 and 50%) and 7-19 weeks from the higher grazing intensities (75 and 100%) across the estuary. Increased grazing intensity led to not only longer recovery times (RTs), but also greater variability in the RT among experimental locations. The RT from the higher grazing intensities at one location in particular was more than double other locations. Seagrass recovery was through vegetative mechanisms and not through sexual reproduction. There was a significant grazing treatment effect on seagrass meadow characteristics, particularly belowground biomass which had not recovered 3 months following grazing. As the pressure of climate change on estuarine environments increases, these quantified RTs for seagrass provide a baseline for understanding grazing pressure as a singular disturbance. Future work can now examine how grazing and other potentially interacting pressures in our changing climate could impact seagrass recovery even further.

Impacts of land-use change and urban development on carbon sequestration in tropical seagrass meadow sediments.

Seagrass meadows store significant carbon stocks at a global scale, but land-use change and other anthropogenic activities can alter the natural process of organic carbon (Corg) accumulation. Here, we assessed the carbon accumulation history of two seagrass meadows in Zanzibar (Tanzania) that have experienced different degrees of disturbance. The meadow at Stone Town has been highly exposed to urban development during the 20th century, while the Mbweni meadow is located in an area with relatively low impacts but historical clearing of adjacent mangroves. The results showed that the two sites had similar sedimentary Corg accumulation rates (22-25 g m-2 yr-1) since the 1940s, while during the last two decades (∼1998 until 2018) they exhibited 24-30% higher accumulation of Corg, which was linked to shifts in Corg sources. The increase in the δ13C isotopic signature of sedimentary Corg (towards a higher seagrass contribution) at the Stone Town site since 1998 points to improved seagrass meadow conditions and Corg accumulation capacity of the meadow after the relocation of a major sewage outlet in the mid-1990s. In contrast, the decrease in the δ13C signatures of sedimentary Corg in the Mbweni meadow since the early 2010s was likely linked to increased Corg run-off of mangrove/terrestrial material following mangrove deforestation. This study exemplifies two different pathways by which land-based human activities can alter the carbon storage capacity of seagrass meadows (i.e. sewage waste management and mangrove deforestation) and showcases opportunities for management of vegetated coastal Corg sinks.

Impact of seagrass establishment, industrialization and coastal infrastructure on seagrass biogeochemical sinks.

The study of a Posidonia sinuosa sedimentary archive has delivered a millenary record of environmental change in Cockburn Sound (Western Australia). Ecosystem change is a major environmental problem challenging sustainable coastal development worldwide, and this study shows baseline trends and shifts in ecological processes in coastal ecosystems under environmental stress. The concentrations and fluxes of biogeochemical elements over the last 3,500 years indicate that important changes in ecosystem dynamics occurred over the last 1,000 years, in particular after ~1900's, probably related to establishment of seagrass meadows in the area and to local and regional human activities (industry and coastal development), respectively. The establishment of seagrasses ~1,000 years ago in the area of study is supported by the appearance of Posidonia fibres from ~40 cm soil depth until the core top, higher δ13C values indicating a larger contribution of seagrass-matter to the soil organic carbon pool, and increased concentration of fine sediments driven by the effect of seagrass canopy in enhancing sedimentation. The comparison of organic carbon, nutrients and metal concentrations and fluxes between pre- and post-establishment of seagrasses shows that seagrass establishment resulted in up to 9-fold increase in the soil biogeochemical sink. In ~1900's, shifts in the concentrations of metals, carbonates, organic carbon, sediment grain size, and δ13C and δ15N values of the organic matter were detected, demonstrating an alteration in seagrass ecosystem functioning following the onset of European settlement. Anthropogenic activities, and in particular the construction of a causeway in 1970's, enhanced seagrass soil organic carbon and metal accumulation rates by 36- and 39-fold, respectively, showing that human-made structures can enhance the biogeochemical sink capacity of seagrasses. Here we reconstruct the impact of human activities on seagrass ecosystem dynamics and blue carbon, which can inform local management of Cockburn Sound and seagrass conservation for climate change mitigation and adaptation.

Seagrass losses since mid-20th century fuelled CO2 emissions from soil carbon stocks.

Seagrass meadows store globally significant organic carbon (Corg ) stocks which, if disturbed, can lead to CO2 emissions, contributing to climate change. Eutrophication and thermal stress continue to be a major cause of seagrass decline worldwide, but the associated CO2 emissions remain poorly understood. This study presents comprehensive estimates of seagrass soil Corg erosion following eutrophication-driven seagrass loss in Cockburn Sound (23 km2 between 1960s and 1990s) and identifies the main drivers. We estimate that shallow seagrass meadows (<5 m depth) had significantly higher Corg stocks in 50 cm thick soils (4.5 ± 0.7 kg Corg /m2 ) than previously vegetated counterparts (0.5 ± 0.1 kg Corg /m2 ). In deeper areas (>5 m), however, soil Corg stocks in seagrass and bare but previously vegetated areas were not significantly different (2.6 ± 0.3 and 3.0 ± 0.6 kg Corg /m2 , respectively). The soil Corg sequestration capacity prevailed in shallow and deep vegetated areas (55 ± 11 and 21 ± 7 g Corg  m-2  year-1 , respectively), but was lost in bare areas. We identified that seagrass canopy loss alone does not necessarily drive changes in soil Corg but, when combined with high hydrodynamic energy, significant erosion occurred. Our estimates point at ~0.20 m/s as the critical shear velocity threshold causing soil Corg erosion. We estimate, from field studies and satellite imagery, that soil Corg erosion (within the top 50 cm) following seagrass loss likely resulted in cumulative emissions of 0.06-0.14 Tg CO2-eq over the last 40 years in Cockburn Sound. We estimated that indirect impacts (i.e. eutrophication, thermal stress and light stress) causing the loss of ~161,150 ha of seagrasses in Australia, likely resulted in the release of 11-21 Tg CO2 -eq since the 1950s, increasing cumulative CO2 emissions from land-use change in Australia by 1.1%-2.3% per annum. The patterns described serve as a baseline to estimate potential CO2 emissions following disturbance of seagrass meadows.

Factors regulating primary producers' assemblages in Posidonia oceanica (L.) Delile ecosystems over the past 1800 years.

Posidonia oceanica (L.) Delile meadows are highly productive coastal marine ecosystems that provide multiple ecosystem services. The seagrass is not always the major contributor to total primary production, however, little is known about long-term changes in the composition of primary producers within seagrass meadows. Understanding compositional shifts within the community of primary producers is crucial to evaluate how climate and anthropogenic change affect the functioning of seagrass ecosystems. Here we analysed marker pigment composition in seagrass cores from two bays of the Cabrera Island (Balearic Islands, Spain) to asses long-term changes in phototrophic community composition and production in seagrass meadows, and identify the environmental factors triggering those changes. The proxy dataset was explored using principal component analyses (PCA): one including the pigment dataset to look for associations between producers' groups, and another one combining the pigment dataset with plausible local and global regulatory factors to assess the environmental drivers of change. Analyses of characteristic pigments and morphological fossils (cysts) showed that the abundance of dinoflagellates increased over the last 150-300 years, coeval with a rise in solar irradiance and air temperature. When compared among embayments, pigments from cyanobacteria predominated in seagrass meadows located at Es Port, a sheltered bay receiving higher terrestrial runoff; whereas pigments from diatoms, seagrasses and rodophytes were more common at Santa Maria, an exposed bay with clearer waters. Water depth also played a role in controlling the phototrophic community composition, with greater abundance of diatoms in the shallowest waters (<5 m). Overall, our results suggested that historical and spatial variation in seagrass meadows' phototrophic community composition was influenced by the interaction between local factors (catchment-bay characteristics) and global climate processes (energy influx). Together these patterns forecast how marine primary producers and seagrass ecosystem structure may respond to future global warming.

Author Correction: The future of Blue Carbon science.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Australian vegetated coastal ecosystems as global hotspots for climate change mitigation.

Policies aiming to preserve vegetated coastal ecosystems (VCE; tidal marshes, mangroves and seagrasses) to mitigate greenhouse gas emissions require national assessments of blue carbon resources. Here, we present organic carbon (C) storage in VCE across Australian climate regions and estimate potential annual CO2 emission benefits of VCE conservation and restoration. Australia contributes 5-11% of the C stored in VCE globally (70-185 Tg C in aboveground biomass, and 1,055-1,540 Tg C in the upper 1 m of soils). Potential CO2 emissions from current VCE losses are estimated at 2.1-3.1 Tg CO2-e yr-1, increasing annual CO2 emissions from land use change in Australia by 12-21%. This assessment, the most comprehensive for any nation to-date, demonstrates the potential of conservation and restoration of VCE to underpin national policy development for reducing greenhouse gas emissions.

The future of Blue Carbon science.

The term Blue Carbon (BC) was first coined a decade ago to describe the disproportionately large contribution of coastal vegetated ecosystems to global carbon sequestration. The role of BC in climate change mitigation and adaptation has now reached international prominence. To help prioritise future research, we assembled leading experts in the field to agree upon the top-ten pending questions in BC science. Understanding how climate change affects carbon accumulation in mature BC ecosystems and during their restoration was a high priority. Controversial questions included the role of carbonate and macroalgae in BC cycling, and the degree to which greenhouse gases are released following disturbance of BC ecosystems. Scientists seek improved precision of the extent of BC ecosystems; techniques to determine BC provenance; understanding of the factors that influence sequestration in BC ecosystems, with the corresponding value of BC; and the management actions that are effective in enhancing this value. Overall this overview provides a comprehensive road map for the coming decades on future research in BC science.

Habitat characteristics provide insights of carbon storage in seagrass meadows.

Seagrass meadows provide multiple ecosystem services, yet they are among the most threatened ecosystems on earth. Because of their role as carbon sinks, protection and restoration of seagrass meadows contribute to climate change mitigation. Blue Carbon strategies aim to enhance CO2 sequestration and avoid greenhouse gasses emissions through the management of coastal vegetated ecosystems, including seagrass meadows. The implementation of Blue Carbon strategies requires a good understanding of the habitat characteristics that influence Corg sequestration. Here, we review the existing knowledge on Blue Carbon research in seagrass meadows to identify the key habitat characteristics that influence Corg sequestration in seagrass meadows, those factors that threaten this function and those with unclear effects. We demonstrate that not all seagrass habitats have the same potential, identify research priorities and describe the implications of the results found for the implementation and development of efficient Blue Carbon strategies based on seagrass meadows.

Disturbance Is an Important Driver of Clonal Richness in Tropical Seagrasses.

Clonality is common in many aquatic plant species, including seagrasses, where populations are maintained through a combination of asexual and sexual reproduction. One common measure used to describe the clonal structure of populations is clonal richness. Clonal richness is strongly dependent on the biological characteristics of the species, and how these interact with the environment but can also reflect evolutionary scale processes especially at the edge of species ranges. However, little is known about the spatial patterns and drivers of clonal richness in tropical seagrasses. This study assessed the spatial patterns of clonal richness in meadows of three tropical seagrass species, Thalassia hemprichii, Halodule uninervis, and Halophila ovalis, spanning a range of life-history strategies and spatial scales (2.5-4,711 km) in Indonesia and NW Australia. We further investigated the drivers of clonal richness using general additive mixed models for two of the species, H. uninervis and H. ovalis, over 8° latitude. No significant patterns were observed in clonal richness with latitude, yet disturbance combined with sea surface temperature strongly predicted spatial patterns of clonal richness. Sites with a high probability of cyclone disturbance had low clonal richness, whereas an intermediate probability of cyclone disturbance and the presence of dugong grazing combined with higher sea surface temperatures resulted in higher levels of clonal richness. We propose potential mechanisms for these patterns related to the recruitment and mortality rates of individuals as well as reproductive effort. Under a changing climate, increased severity of tropical cyclones and the decline in populations of mega-grazers have the potential to reduce clonal richness leading to less genetically diverse populations.

Response of the seagrass Halophila ovalis to altered light quality in a simulated dredge plume.

Seagrass meadows are globally threatened, largely through activities that reduce light quantity (photosynthetic photon flux density) such as dredging. However, these activities can simultaneously alter the spectral quality of light. Previous studies showed that Halophila ovalis seagrass productivity is reduced under monochromatic yellow/green light, wavelengths associated with dredge plumes, but it is unclear how they respond to spectra produced by real dredging projects. We simultaneously subjected adult H. ovalis plants to altered light quality and quantity simulating a real commercial dredging operation (15mgL-1 TSS, 50 and 200µmol photonsm-2s-1). There was a significant effect of reduced light quantity on physiological and morphological variables and a significant effect of light quality on the pigment antheraxanthin. The lack of effect of light quality on growth indicates that while seagrass are sensitive to changes in light quality, natural- and anthropogenic-driven changes may not always be sufficient to produce strong effects on H. ovalis.

Identifying knowledge gaps in seagrass research and management: An Australian perspective.

Seagrass species form important marine and estuarine habitats providing valuable ecosystem services and functions. Coastal zones that are increasingly impacted by anthropogenic development have experienced substantial declines in seagrass abundance around the world. Australia, which has some of the world's largest seagrass meadows and is home to over half of the known species, is not immune to these losses. In 1999 a review of seagrass ecosystems knowledge was conducted in Australia and strategic research priorities were developed to provide research direction for future studies and management. Subsequent rapid evolution of seagrass research and scientific methods has led to more than 70% of peer reviewed seagrass literature being produced since that time. A workshop was held as part of the Australian Marine Sciences Association conference in July 2015 in Geelong, Victoria, to update and redefine strategic priorities in seagrass research. Participants identified 40 research questions from 10 research fields (taxonomy and systematics, physiology, population biology, sediment biogeochemistry and microbiology, ecosystem function, faunal habitats, threats, rehabilitation and restoration, mapping and monitoring, management tools) as priorities for future research on Australian seagrasses. Progress in research will rely on advances in areas such as remote sensing, genomic tools, microsensors, computer modeling, and statistical analyses. A more interdisciplinary approach will be needed to facilitate greater understanding of the complex interactions among seagrasses and their environment.

Historical processes and contemporary ocean currents drive genetic structure in the seagrass Thalassia hemprichii in the Indo-Australian Archipelago.

Understanding spatial patterns of gene flow and genetic structure is essential for the conservation of marine ecosystems. Contemporary ocean currents and historical isolation due to Pleistocene sea level fluctuations have been predicted to influence the genetic structure in marine populations. In the Indo-Australian Archipelago (IAA), the world's hotspot of marine biodiversity, seagrasses are a vital component but population genetic information is very limited. Here, we reconstructed the phylogeography of the seagrass Thalassia hemprichii in the IAA based on single nucleotide polymorphisms (SNPs) and then characterized the genetic structure based on a panel of 16 microsatellite markers. We further examined the relative importance of historical isolation and contemporary ocean currents in driving the patterns of genetic structure. Results from SNPs revealed three population groups: eastern Indonesia, western Indonesia (Sunda Shelf) and Indian Ocean; while the microsatellites supported five population groups (eastern Indonesia, Sunda Shelf, Lesser Sunda, Western Australia and Indian Ocean). Both SNPs and microsatellites showed asymmetrical gene flow among population groups with a trend of southwestward migration from eastern Indonesia. Genetic diversity was generally higher in eastern Indonesia and decreased southwestward. The pattern of genetic structure and connectivity is attributed partly to the Pleistocene sea level fluctuations modified to a smaller level by contemporary ocean currents.

Short-term Responses of Posidonia australis to Changes in Light Quality.

Seagrass meadows are highly productive ecosystems that provide ecosystem services to the coastal zone but are declining globally, particularly due to anthropogenic activities that reduce the quantity of light reaching seagrasses, such as dredging, river discharge and eutrophication. Light quality (the spectral composition of the light) is also altered by these anthropogenic stressors as the differential attenuation of wavelengths of light is caused by materials within the water column. This study addressed the effect of altered light quality on different life-history stages of the seagrass Posidonia australis, a persistent, habitat-forming species in Australia. Aquarium-based experiments were conducted to determine how adult shoots and seedlings respond to blue (peak λ = 451 nm); green (peak λ = 522 nm); yellow (peak λ = 596 nm) and red (peak λ = 673 nm) wavelengths with a control of full-spectrum light (λ = 400 - 700 nm, at 200 µmol photons m-2 s-1). Posidonia australis adults did not respond to changes in light quality relative to full-spectrum light, demonstrating a capacity to obtain enough photons from a range of wavelengths across the visible spectrum to maintain short-term growth at high irradiances. Posidonia australis seedlings (<4 months old) grown in blue light showed a significant increase in xanthophyll concentrations when compared to plants grown in full-spectrum, demonstrating a pigment acclimation response to blue light. These results differed significantly from negative responses to changes in light quality recently described for Halophila ovalis, a colonizing seagrass species. Persistent seagrasses such as P. australis, appear to be better at tolerating short-term changes in light quality compared to colonizing species when sufficient PPFD is present.

Coastal connectivity and spatial subsidy from a microbial perspective.

The transfer of organic material from one coastal environment to another can increase production in recipient habitats in a process known as spatial subsidy. Microorganisms drive the generation, transformation, and uptake of organic material in shallow coastal environments, but their significance in connecting coastal habitats through spatial subsidies has received limited attention. We address this by presenting a conceptual model of coastal connectivity that focuses on the flow of microbially mediated organic material in key coastal habitats. Our model suggests that it is not the difference in generation rates of organic material between coastal habitats but the amount of organic material assimilated into microbial biomass and respiration that determines the amount of material that can be exported from one coastal environment to another. Further, the flow of organic material across coastal habitats is sensitive to environmental change as this can alter microbial remineralization and respiration rates. Our model highlights microorganisms as an integral part of coastal connectivity and emphasizes the importance of including a microbial perspective in coastal connectivity studies.

Utilization of carbon substrates by heterotrophic bacteria through vertical sediment profiles in coastal and estuarine seagrass meadows.

Coastal vegetated ecosystems play an important role in carbon cycling and bacterial communities inhabiting coastal sediments are responsible for the remineralization and processing of organic carbon (OC). We collected 1 m-long sediment cores in Posidonia seagrass meadows from coastal and estuarine sites in Australia that differed in their sedimentary organic and inorganic carbon, nitrogen and mud contents. The metabolic diversity of sediment heterotrophic bacterial communities was characterized at different sediment depths, based on the utilization pattern of 31 individual carbon substrates using Biolog EcoPlatesTM . High metabolic diversity was recorded at both sites, but the carbon substrate utilization rates and the use of carbohydrates were higher at the coastal site compared to the estuarine site. The heterotrophic bacterial community in the coastal sediment appeared to metabolize a more diverse OC pool compared to the estuarine site, which might partly explain the differences in OC storage among the seagrass habitats studied. The Biolog EcoPlatesTM provided a useful tool for characterising the sediment heterotrophic bacterial communities in the meadows and sediment characteristics and biochemical composition of the organic matter played an important role in shaping heterotrophic bacterial communities and their carbon utilization rates, potentially affecting carbon accumulation and preservation within seagrass sediments.

Accelerating Tropicalization and the Transformation of Temperate Seagrass Meadows.

Climate-driven changes are altering production and functioning of biotic assemblages in terrestrial and aquatic environments. In temperate coastal waters, rising sea temperatures, warm water anomalies and poleward shifts in the distribution of tropical herbivores have had a detrimental effect on algal forests. We develop generalized scenarios of this form of tropicalization and its potential effects on the structure and functioning of globally significant and threatened seagrass ecosystems, through poleward shifts in tropical seagrasses and herbivores. Initially, we expect tropical herbivorous fishes to establish in temperate seagrass meadows, followed later by megafauna. Tropical seagrasses are likely to establish later, delayed by more limited dispersal abilities. Ultimately, food webs are likely to shift from primarily seagrass-detritus to more direct-consumption-based systems, thereby affecting a range of important ecosystem services that seagrasses provide, including their nursery habitat role for fishery species, carbon sequestration, and the provision of organic matter to other ecosystems in temperate regions.

Reconstruction of centennial-scale fluxes of chemical elements in the Australian coastal environment using seagrass archives.

The study of a Posidonia australis sedimentary archive has provided a record of changes in element concentrations (Al, Fe, Mn, Pb, Zn, Cr, Cd, Co, As, Cu, Ni and S) over the last 3000 years in the Australian marine environment. Human-derived contamination in Oyster Harbor (SW Australia) started ~100 years ago (AD ~1900) and exponentially increased until present. This appears to be related to European colonization of Australia and the subsequent impact of human activities, namely mining, coal and metal production, and extensive agriculture. Two contamination periods of different magnitude have been identified: Expansion period (EXP, AD ~1900-1970) and Establishment period (EST, AD ~1970 to present). Enrichments of chemical elements with respect to baseline concentrations (in samples older than ~115 cal years BP) were found for all elements studied in both periods, except for Ni, As and S. The highest enrichment factors were obtained for the EST period (ranging from 1.3-fold increase in Cu to 7.2-fold in Zn concentrations) compared to the EXP period (1.1-fold increase for Cu and Cr to 2.4-fold increase for Pb). Zinc, Pb, Mn and Co concentrations during both periods were 2- to 7-fold higher than baseline levels. This study demonstrates the value of Posidonia mats as long-term archives of element concentrations and trends in coastal ecosystems. We also provide preliminary evidence on the potential for Posidonia meadows to act as significant long-term biogeochemical sinks of chemical elements.

Phenological decoupling of mortality from wave forcing in kelp beds.

Kelps often live in a harsh hydrodynamic environment where wave-driven dislodgement of individuals can alter the biodiversity and functioning of reef systems, and increase production in coastal ecosystems adjacent to reefs. The current paradigm is that winter storms tear kelps from reefs once hydrodynamic forces exceed attachment or tissue strength--a threshold response that implies a pulsed relationship between wave forces and dislodgement. Here, we challenge this understanding by showing how kelp phenology can decouple susceptibility to dislodgement from seasonal patterns in wave forces. We measured kelp dislodgement rates and hydrodynamic forces at nine subtidal reefs over two years (n = 4320 kelps tagged and monitored). Contrary to expectation, we found relatively low and constant dislodgement rates for all reefs (13% +/- 6% [mean per season +/- SD]) in spite of a strong temporal pattern in wave action and extreme water velocities (winter peaks up to 3-4 m/s). A biomechanical model, based on the balance between kelp attachment strength and hydrodynamic drag, demonstrated that severe reduction in individual kelp size toward winter (>50% decrease in biomass for all sites) minimized drag and made the kelps less susceptible to high water velocities, allowing individuals to survive storm velocities over 3-4 m/s. We conclude that the timing of reduced susceptibility to disturbance, through the seasonal reduction of individual kelp biomass that coincides with times of highest water velocities is critical to the dynamics of kelp dislodgement and survival. We propose that phenological processes maintain many kelp beds in a higher degree of population stability and equilibrium with hydrodynamic forces than previously believed.