Quantifying blue carbon stocks and the role of protected areas to conserve coastal wetlands.

Vegetated coastal ecosystems, in particular mangroves, tidal marshes and seagrasses are highly efficient at sequestering and storing carbon, making them valuable assets for climate change mitigation and adaptation. The state of Queensland, in northeastern Australia, contains almost half of the total area of these blue carbon ecosystems in the country, yet there are few detailed regional or state-wide assessments of their total sedimentary organic carbon (SOC) stocks. We compiled existing SOC data and used boosted regression tree models to evaluate the influence of environmental variables in explaining the variability in SOC stocks, and to produce spatially explicit blue carbon estimates. The final models explained 75 % (for mangroves and tidal marshes) and 65 % (for seagrasses) of the variability in SOC stocks. Total SOC stocks in the state of Queensland were estimated at 569 ± 98 Tg C (173 ± 32 Tg C, 232 ± 50 Tg C, and 164 ± 16 Tg C from mangroves, tidal marshes and seagrasses, respectively). Regional predictions for each of Queensland's eleven Natural Resource Management regions revealed that 60 % of the state's SOC stocks occurred within three regions (Cape York, Torres Strait and Southern Gulf Natural Resource Management regions) due to a combination of high values of SOC stocks and large areas of coastal wetlands. Protected areas in Queensland play an important role in conserving SOC assets in Queensland's coastal wetlands. For example, ~19 Tg C within terrestrial protected areas, ~27 Tg C within marine protected areas and ~ 40 Tg C within areas of matters of State Environmental Significance. Using multi-decadal (1987-2020) mapped distributions of mangroves in Queensland; we found that mangrove area increased by approximately 30,000 ha from 1987 to 2020, which led to temporal fluctuations in mangrove plant and SOC stocks. We estimated that plant stocks decreased from ~45 Tg C in 1987 to ~34.2 Tg C in 2020, while SOC stocks remained relatively constant from ~107.9 Tg C in 1987 to 108.0 Tg C in 2020. Considering the level of current protection, emissions from mangrove deforestation are potentially very low; therefore, representing minor opportunities for mangrove blue carbon projects in the region. Our study provides much needed information on current trends in carbon stocks and their conservation in Queensland's coastal wetlands, while also contributing to guide future management actions, including blue carbon restoration projects.

Current and future carbon stocks in coastal wetlands within the Great Barrier Reef catchments.

Australia's Great Barrier Reef (GBR) catchments include some of the world's most intact coastal wetlands comprising diverse mangrove, seagrass and tidal marsh ecosystems. Although these ecosystems are highly efficient at storing carbon in marine sediments, their soil organic carbon (SOC) stocks and the potential changes resulting from climate impacts, including sea level rise are not well understood. For the first time, we estimated SOC stocks and their drivers within the range of coastal wetlands of GBR catchments using boosted regression trees (i.e. a machine learning approach and ensemble method for modelling the relationship between response and explanatory variables) and identified the potential changes in future stocks due to sea level rise. We found levels of SOC stocks of mangrove and seagrass meadows have different drivers, with climatic variables such as temperature, rainfall and solar radiation, showing significant contributions in accounting for variation in SOC stocks in mangroves. In contrast, soil type accounted for most of the variability in seagrass meadows. Total SOC stock in the GBR catchments, including mangroves, seagrass meadows and tidal marshes, is approximately 137 Tg C, which represents 9%-13% of Australia's total SOC stock while encompassing only 4%-6% of the total extent of Australian coastal wetlands. In a global context, this could represent 0.5%-1.4% of global SOC stock. Our study suggests that landward migration due to projected sea level rise has the potential to enhance carbon accumulation with total carbon gains between 0.16 and 0.46 Tg C and provides an opportunity for future restoration to enhance blue carbon.

Ecosystem type drives tea litter decomposition and associated prokaryotic microbiome communities in freshwater and coastal wetlands at a continental scale.

Wetland ecosystems are critical to the regulation of the global carbon cycle, and there is a high demand for data to improve carbon sequestration and emission models and predictions. Decomposition of plant litter is an important component of ecosystem carbon cycling, yet a lack of knowledge on decay rates in wetlands is an impediment to predicting carbon preservation. Here, we aim to fill this knowledge gap by quantifying the decomposition of standardised green and rooibos tea litter over one year within freshwater and coastal wetland soils across four climates in Australia. We also captured changes in the prokaryotic members of the tea-associated microbiome during this process. Ecosystem type drove differences in tea decay rates and prokaryotic microbiome community composition. Decomposition rates were up to 2-fold higher in mangrove and seagrass soils compared to freshwater wetlands and tidal marshes, in part due to greater leaching-related mass loss. For tidal marshes and freshwater wetlands, the warmer climates had 7-16% less mass remaining compared to temperate climates after a year of decomposition. The prokaryotic microbiome community composition was significantly different between substrate types and sampling times within and across ecosystem types. Microbial indicator analyses suggested putative metabolic pathways common across ecosystems were used to breakdown the tea litter, including increased presence of putative methylotrophs and sulphur oxidisers linked to the introduction of oxygen by root in-growth over the incubation period. Structural equation modelling analyses further highlighted the importance of incubation time on tea decomposition and prokaryotic microbiome community succession, particularly for rooibos tea that experienced a greater proportion of mass loss between three and twelve months compared to green tea. These results provide insights into ecosystem-level attributes that affect both the abiotic and biotic controls of belowground wetland carbon turnover at a continental scale, while also highlighting new decay dynamics for tea litter decomposing under longer incubations.

Climate drives the geography of marine consumption by changing predator communities.

The global distribution of primary production and consumption by humans (fisheries) is well-documented, but we have no map linking the central ecological process of consumption within food webs to temperature and other ecological drivers. Using standardized assays that span 105° of latitude on four continents, we show that rates of bait consumption by generalist predators in shallow marine ecosystems are tightly linked to both temperature and the composition of consumer assemblages. Unexpectedly, rates of consumption peaked at midlatitudes (25 to 35°) in both Northern and Southern Hemispheres across both seagrass and unvegetated sediment habitats. This pattern contrasts with terrestrial systems, where biotic interactions reportedly weaken away from the equator, but it parallels an emerging pattern of a subtropical peak in marine biodiversity. The higher consumption at midlatitudes was closely related to the type of consumers present, which explained rates of consumption better than consumer density, biomass, species diversity, or habitat. Indeed, the apparent effect of temperature on consumption was mostly driven by temperature-associated turnover in consumer community composition. Our findings reinforce the key influence of climate warming on altered species composition and highlight its implications for the functioning of Earth's ecosystems.

Green turtle (Chelonia mydas) grazing plot formation creates structural changes in a multi-species Great Barrier Reef seagrass meadow.

The Great Barrier Reef (GBR) contains extensive seagrass meadows with abundant and diverse herbivore populations. Typically, meadows in the region are multi-species and dominated by fast growing opportunistic seagrass species. However, we know little about how herbivores modify these types of seagrass meadows by grazing. We conducted the first megaherbivore exclusion study in the GBR at Green Island (Queensland) to understand how green turtle grazing structures these multi-species tropical seagrass meadows. After excluding green turtles for three months, we found that grazing only impacted seagrasses at one site, where green turtles created a grazing plot by actively feeding on both above and below ground seagrass structures, a rare observation for the species. Within this grazing plot at the end of the experiment, the un-caged control treatments open to grazing had a 60% reduction in both above and below ground biomass, and shoot height was reduced by 75%, but there was no impact of grazing on the seagrass species mix. Our study shows that grazing plot formation by green turtles occurs in GBR fast growing seagrass communities and reduces both above and below ground seagrass biomass, this behaviour may be targeting elevated leaf nutrients, or nutritional content of rhizomes. This study is the first documented case of grazing plot formation by green turtles in the GBR and suggests that grazing pressure has a major influence on seagrass meadow structure.

High variability of Blue Carbon storage in seagrass meadows at the estuary scale.

Seagrass meadows are considered important natural carbon sinks due to their capacity to store organic carbon (Corg) in sediments. However, the spatial heterogeneity of carbon storage in seagrass sediments needs to be better understood to improve accuracy of Blue Carbon assessments, particularly when strong gradients are present. We performed an intensive coring study within a sub-tropical estuary to assess the spatial variability in sedimentary Corg associated with seagrasses, and to identify the key factors promoting this variability. We found a strong spatial pattern within the estuary, from 52.16 mg Corg cm-3 in seagrass meadows in the upper parts, declining to 1.06 mg Corg cm-3 in seagrass meadows at the estuary mouth, despite a general gradient of increasing seagrass cover and seagrass habitat extent in the opposite direction. The sedimentary Corg underneath seagrass meadows came principally from allochthonous (non-seagrass) sources (~70-90 %), while the contribution of seagrasses was low (~10-30 %) throughout the entire estuary. Our results showed that Corg stored in sediments of seagrass meadows can be highly variable within an estuary, attributed largely to accumulation of fine sediments and inputs of allochthonous sources. Local features and the existence of spatial gradients must be considered in Blue Carbon estimates in coastal ecosystems.

The Role of Herbivory in Structuring Tropical Seagrass Ecosystem Service Delivery.

Seagrass meadows support key ecosystem services, via provision of food directly for herbivores, and indirectly to their predators. The importance of herbivores in seagrass meadows has been well-documented, but the links between food webs and ecosystem services in seagrass meadows have not previously been made explicit. Herbivores interact with ecosystem services - including carbon sequestration, cultural values, and coastal protection. Interactions can be positive or negative and depend on a range of factors including the herbivore identity and the grazing type and intensity. There can be unintended consequences from management actions based on a poor understanding of trade-offs that occur with complex seagrass-herbivore interactions. Tropical seagrass meadows support a diversity of grazers spanning the meso-, macro-, and megaherbivore scales. We present a conceptual model to describe how multiple ecosystem services are influenced by herbivore pressure in tropical seagrass meadows. Our model suggests that a balanced ecosystem, incorporating both seagrass and herbivore diversity, is likely to sustain the broadest range of ecosystem services. Our framework suggests the pathway to achieve desired ecosystem services outcomes requires knowledge on four key areas: (1) how size classes of herbivores interact to structure seagrass; (2) desired community and management values; (3) seagrass responses to top-down and bottom-up controls; (4) the pathway from intermediate to final ecosystem services and human benefits. We suggest research should be directed to these areas. Herbivory is a major structuring influence in tropical seagrass systems and needs to be considered for effective management of these critical habitats and their services.

Contrasting impacts of light reduction on sediment biogeochemistry in deep- and shallow-water tropical seagrass assemblages (Green Island, Great Barrier Reef).

Seagrass meadows increasingly face reduced light availability as a consequence of coastal development, eutrophication, and climate-driven increases in rainfall leading to turbidity plumes. We examined the impact of reduced light on above-ground seagrass biomass and sediment biogeochemistry in tropical shallow- (∼2 m) and deep-water (∼17 m) seagrass meadows (Green Island, Australia). Artificial shading (transmitting ∼10-25% of incident solar irradiance) was applied to the shallow- and deep-water sites for up to two weeks. While above-ground biomass was unchanged, higher diffusive O2 uptake (DOU) rates, lower O2 penetration depths, and higher volume-specific O2 consumption (R) rates were found in seagrass-vegetated sediments as compared to adjacent bare sand (control) areas at the shallow-water sites. In contrast, deep-water sediment characteristics did not differ between bare sand and vegetated sites. At the vegetated shallow-water site, shading resulted in significantly lower hydrogen sulphide (H2S) levels in the sediment. No shading effects were found on sediment biogeochemistry at the deep-water site. Overall, our results show that the sediment biogeochemistry of shallow-water (Halodule uninervis, Syringodium isoetifolium, Cymodocea rotundata and C. serrulata) and deep-water (Halophila decipiens) seagrass meadows with different species differ in response to reduced light. The light-driven dynamics of the sediment biogeochemistry at the shallow-water site could suggest the presence of a microbial consortium, which might be stimulated by photosynthetically produced exudates from the seagrass, which becomes limited due to lower seagrass photosynthesis under shaded conditions.

Blue Carbon stocks of Great Barrier Reef deep-water seagrasses.

Shallow-water seagrasses capture and store globally significant quantities of organic carbon (OC), often referred to as 'Blue Carbon'; however, data are lacking on the importance of deep-water (greater than 15 m) seagrasses as Blue Carbon sinks. We compared OC stocks from deep-, mid- and shallow-water seagrasses at Lizard Island within the Great Barrier Reef (GBR) lagoon. We found deep-water seagrass ( Halophila species) contained similar levels of OC to shallow-water species (e.g. Halodule uninervis) (0.64 ± 0.08% and 0.9 ± 0.1 mg C cm-3, 0.87 ± 0.19% and 1.3 ± 0.3 mg C cm-3, respectively), despite being much sparser and smaller in stature. Deep-water seagrass sediments contained significantly higher levels (approx. ninefold) of OC than surrounding bare areas. Inorganic carbon (CaCO3) levels were relatively high in deep-water seagrass sediments (8.2 ± 0.4%) and, if precipitated from epiphytes within the meadow, could offset the potential CO2-sink capacity of these meadows. The δ13C signatures of sediment samples varied among depths and habitats (-10.9 and -17.0), reflecting contributions from autochthonous and allochthonous sources. If the OC stocks reported in this study are similar to deep-water Halophila meadows elsewhere within the GBR lagoon (total area 31 000 km2), then OC bound within this system is roughly estimated at 27.4 million tonnes.

Timing anthropogenic stressors to mitigate their impact on marine ecosystem resilience.

Better mitigation of anthropogenic stressors on marine ecosystems is urgently needed to address increasing biodiversity losses worldwide. We explore opportunities for stressor mitigation using whole-of-systems modelling of ecological resilience, accounting for complex interactions between stressors, their timing and duration, background environmental conditions and biological processes. We then search for ecological windows, times when stressors minimally impact ecological resilience, defined here as risk, recovery and resistance. We show for 28 globally distributed seagrass meadows that stressor scheduling that exploits ecological windows for dredging campaigns can achieve up to a fourfold reduction in recovery time and 35% reduction in extinction risk. Although the timing and length of windows vary among sites to some degree, global trends indicate favourable windows in autumn and winter. Our results demonstrate that resilience is dynamic with respect to space, time and stressors, varying most strongly with: (i) the life history of the seagrass genus and (ii) the duration and timing of the impacting stress.

Long distance biotic dispersal of tropical seagrass seeds by marine mega-herbivores.

Terrestrial plants use an array of animals as vectors for dispersal, however little is known of biotic dispersal of marine angiosperms such as seagrasses. Our study in the Great Barrier Reef confirms for the first time that dugongs (Dugong dugon) and green sea turtles (Chelonia mydas) assist seagrass dispersal. We demonstrate that these marine mega-herbivores consume and pass in faecal matter viable seeds for at least three seagrass species (Zostera muelleri, Halodule uninervis and Halophila decipiens). One to two seagrass seeds per g DW of faecal matter were found during the peak of the seagrass reproductive season (September to December), with viability on excretion of 9.13% ± 4.61% (SE). Using population estimates for these mega-herbivores, and data on digestion time (hrs), average daily movement (km h) and numbers of viable seagrass seeds excreted (per g DW), we calculated potential seagrass seed dispersal distances. Dugongs and green sea turtle populations within this region can disperse >500,000 viable seagrass seeds daily, with a maximum dispersal distance of approximately 650 km. Biotic dispersal of tropical seagrass seeds by dugongs and green sea turtles provides a large-scale mechanism that enhances connectivity among seagrass meadows, and aids in resilience and recovery of these coastal habitats.

Does Biodiversity-Ecosystem Function Literature Neglect Tropical Ecosystems?

Current evidence suggests that there is a positive relationship between biodiversity and ecosystem functioning, but few studies have addressed tropical ecosystems where the highest levels of biodiversity occur. We develop two hypotheses for the implications of generalizing from temperate studies to tropical ecosystems, and discuss the need for more tropical research.

Seed germination in a southern Australian temperate seagrass.

In a series of experiments, seeds from a temperate seagrass species, Zostera nigricaulis collected in Port Phillip Bay, Victoria, Australia were exposed to a range of salinities (20 PSU pulse/no pulse, 25 PSU, 30 PSU, 35 PSU), temperatures (13 °C, 17 °C, 22 °C), burial depths (0 cm, 1 cm, 2 cm) and site specific sediment characteristics (fine, medium, coarse) to quantify their impacts on germination rate and maximum overall germination. In southern Australia the seagrass Z. nigricaulis is a common subtidal species; however, little is known about the factors that affect seed germination which is a potential limiting factor in meadow resilience to natural and anthropogenic disturbances. Overall seed germination was low (<20%) with germination decreasing to <10% when seeds were placed in the sediment. When germination of Z. nigricaulis seeds was observed, it was enhanced (greater overall germination and shorter time to germination) when seeds were exposed to a 20 PSU pulse for 24 h, maintained at salinity of 25 PSU, temperatures <13 °C, in sediments with fine or medium grain sand and buried at a depth of <1 cm. These results indicate that germination of Z. nigricaulis seeds under in situ conditions may be seasonally limited by temperatures in southern Australia. Seed germination may be further restricted by salinity as freshwater pulses reaching 20 PSU are typically only observed in Port Phillip Bay following large scale rainfall events. As a result, these populations may be particularly susceptible to disturbance with only a seasonally limited capacity for recovery.

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.

Spatial variation in reproductive effort of a southern Australian seagrass.

In marine environments characterised by habitat-forming plants, the relative allocation of resources into vegetative growth and flowering is an important indicator of plant condition and hence ecosystem health. In addition, the production and abundance of seeds can give clues to local resilience. Flowering density, seed bank, biomass and epiphyte levels were recorded for the temperate seagrass Zostera nigricaulis in Port Phillip Bay, south east Australia at 14 sites chosen to represent several regions with different physicochemical conditions. Strong regional differences were found within the large bay. Spathe and seed density were very low in the north of the bay (3 sites), low in the centre of the bay (2 sites) intermediate in the Outer Geelong Arm (2 sites), high in Swan Bay (2 sites) and very high in the Inner Geelong Arm (3 sites). In the south (2 sites) seed density was low and spathe density was high. These regional patterns were largely consistent for the 5 sites sampled over the three year period. Timing of flowering was consistent across sites, occurring from August until December with peak production in October, except during the third year of monitoring when overall densities were lower and peaked in November. Seagrass biomass, epiphyte load, canopy height and stem density showed few consistent spatial and temporal patterns. Variation in spathe and seed density and morphology across Port Phillip Bay reflects varying environmental conditions and suggests that northern sites may be restricted in their ability to recover from disturbance through sexual reproduction. In contrast, sites in the west and south of the bay have greater potential to recover from disturbances due to a larger seed bank and these sites could act as source populations for sites where seed production is low.

Variability of sedimentary organic carbon in patchy seagrass landscapes.

Seagrass ecosystems, considered among the most efficient carbon sinks worldwide, encompass a wide variety of spatial configurations in the coastal landscape. Here we evaluated the influence of the spatial configuration of seagrass meadows at small scales (metres) on carbon storage in seagrass sediments. We intensively sampled carbon stocks and other geochemical properties (δ(13)C, particle size, depositional fluxes) across seagrass-sand edges in a Zostera muelleri patchy seagrass landscape. Carbon stocks were significantly higher (ca. 20%) inside seagrass patches than at seagrass-sand edges and bare sediments. Deposition was similar among all positions and most of the carbon was from allochthonous sources. Patch level attributes (e.g. edge distance) represent important determinants of the spatial heterogeneity of carbon stocks within seagrass ecosystems. Our findings indicate that carbon stocks of seagrass areas have likely been overestimated by not considering the influence of meadow landscapes, and have important relevance for the design of seagrass carbon stock assessments.

Dynamics of a deep-water seagrass population on the Great Barrier Reef: annual occurrence and response to a major dredging program.

Global seagrass research efforts have focused on shallow coastal and estuarine seagrass populations where alarming declines have been recorded. Comparatively little is known about the dynamics of deep-water seagrasses despite evidence that they form extensive meadows in some parts of the world. Deep-water seagrasses are subject to similar anthropogenic threats as shallow meadows, particularly along the Great Barrier Reef lagoon where they occur close to major population centres. We examine the dynamics of a deep-water seagrass population in the GBR over an 8 year period during which time a major capital dredging project occurred. Seasonal and inter-annual changes in seagrasses were assessed as well as the impact of dredging. The seagrass population was found to occur annually, generally present between July and December each year. Extensive and persistent turbid plumes from a large dredging program over an 8 month period resulted in a failure of the seagrasses to establish in 2006, however recruitment occurred the following year and the regular annual cycle was re-established. Results show that despite considerable inter annual variability, deep-water seagrasses had a regular annual pattern of occurrence, low resistance to reduced water quality but a capacity for rapid recolonisation on the cessation of impacts.

Resilience of Zostera muelleri seagrass to small-scale disturbances: the relative importance of asexual versus sexual recovery.

Resilience is the ability of an ecosystem to recover from disturbance without loss of essential function. Seagrass ecosystems are key marine and estuarine habitats that are under threat from a variety of natural and anthropogenic disturbances. The ability of these ecosystems to recovery from disturbance will to a large extent depend on the internsity and scale of the disturbance, and the relative importance of sexual versus asexual reproduction within populations. Here, we investigated the resilience of Zostera muelleri seagrass (Syn. Zostera capricorni) to small-scale disturbances at four locations in Lake Macquarie - Australia's largest coastal lake - and monitored recovery over a 65-week period. Resilience of Z. muelleri varied significantly with disturbance intensity; Z. muelleri recovered rapidly (within 2 weeks) from low-intensity disturbance (shoot loss), and rates of recovery appeared related to initial shoot length. Recovery via rhizome encroachment (asexual regeneration) from high-intensity disturbance (loss of entire plant) varied among locations, ranging from 18-35 weeks, whereas the ability to recover was apparently lost (at least within the time frame of this study) when recovery depended on sexual regeneration, suggesting that seeds do not provide a mechanism of recovery against intense small-scale disturbances. The lack of sexual recruits into disturbed sites is surprising as our initial surveys of genotypic diversity (using nine polymorphic microsatellite loci) at these location indicate that populations are maintained by a mix of sexual and asexual reproduction (genotypic diversity [R] varied from 0.24 to 0.44), and populations consisted of a mosaic of genotypes with on average 3.6 unique multilocus genotypes per 300 mm diameter plot. We therefore conclude that Z. muelleri populations within Lake Macquarie rely on clonal growth to recover from small-scale disturbances and that ongoing sexual recruitment by seeds into established seagrass beds (as opposed to bare areas arising from disturbance) must be the mechanism responsible for maintaining the observed mixed genetic composition of Z. muelleri seagrass meadows.