Spatial variation in population genomic responses to over a century of anthropogenic change within a tidal marsh songbird.

Combating the current biodiversity crisis requires the accurate documentation of population responses to human-induced ecological change. However, our ability to pinpoint population responses to human activities is often limited to the analysis of populations studied well after the fact. Museum collections preserve a record of population responses to anthropogenic change that can provide critical baseline data on patterns of genetic diversity, connectivity, and population structure prior to the onset of human perturbation. Here, we leverage a spatially replicated time series of specimens to document population genomic responses to the destruction of nearly 90% of coastal habitats occupied by the Savannah sparrow (Passerculus sandwichensis) in California. We sequenced 219 sparrows collected from 1889 to 2017 across the state of California using an exome capture approach. Spatial-temporal analyses of genetic diversity found that the amount of habitat lost was not predictive of genetic diversity loss. Sparrow populations from southern California historically exhibited lower levels of genetic diversity and experienced the most significant temporal declines in genetic diversity. Despite experiencing the greatest levels of habitat loss, we found that genetic diversity in the San Francisco Bay area remained relatively high. This was potentially related to an observed increase in gene flow into the Bay Area from other populations. While gene flow may have minimized genetic diversity declines, we also found that immigration from inland freshwater-adapted populations into tidal marsh populations led to the erosion of divergence at loci associated with tidal marsh adaptation. Shifting patterns of gene flow through time in response to habitat loss may thus contribute to negative fitness consequences and outbreeding depression. Together, our results underscore the importance of tracing the genomic trajectories of multiple populations over time to address issues of fundamental conservation concern.

Reciprocal facilitation between ants and small mammals in tidal marshes.

The role of facilitation in shaping natural communities has primarily been studied in the context of plant assemblages, while its relevance for mobile animals remains less understood. Our study investigates whether reciprocal interspecific facilitation may exist between fire ants (Solenopsis richteri) and cavies (Cavia aperea), two mobile animals, in the SW Atlantic coast brackish marshes. Field samples showed a spatial association between ant mounds and cavies, and that ants prefer to use cavy runways for movement within the marsh. Through experiments involving transplanting the dominant plant, cordgrass (Spartina densiflora), and manipulating cavy presence in areas with and without ant mounds, we observed that cavies forage extensively (and defecate more) near ant mounds. The ants actively remove cavy droppings in their mound vicinity. These ant activities and interactions with cavy droppings led to reduced moisture and organic content while increasing nitrate and phosphate levels in marsh sediment. Consequently, this enhanced plant growth, indirectly facilitating the cavies, which preferred consuming vegetation near ant mounds. These cascading indirect effects persisted over time; even four months after cavies left the marshes, transplanted plants near ant mounds remained larger and exhibited more leaf senescence when exposed to cavy herbivory. Therefore, the networks of positive interactions appear to generate simultaneous selection among species (populations), promoting coexistence within the community. Although complex, these reciprocal facilitative effects among mobile animals may be more common than currently believed and should be further studied to gain a better understanding of the underlying mechanisms driving species coexistence in natural communities.

Response of macrophyte litter decomposition in global blue carbon ecosystems to climate change.

Blue carbon ecosystems (BCEs) are important nature-based solutions for climate change-mitigation. However, current debates question the reliability and contribution of BCEs under future climatic-scenarios. The answer to this question depends on ecosystem processes driving carbon-sequestration and -storage, such as primary production and decomposition, and their future rates. We performed a global meta-analysis on litter decomposition rate constants (k) in BCEs and predicted changes in carbon release from 309 studies. The relationships between k and climatic factors were examined by extracting remote-sensing data on air temperature, sea-surface temperature, and precipitation aligning to the decomposition time of each experiment. We constructed global numerical models of litter decomposition to forecast k and carbon release under different scenarios. The current k averages at 27 ± 3 × 10-2 day-1 for macroalgae were higher than for seagrasses (1.7 ± 0.2 × 10-2 day-1 ), mangroves (1.6 ± 0.1 × 10-2 day-1 ) and tidal marshes (5.9 ± 0.5 × 10-3 day-1 ). Macrophyte k increased with both air temperature and precipitation in intertidal BCEs and with sea surface temperature for subtidal seagrasses. Above a temperature threshold for vascular plant litter at ~25°C and ~20°C for macroalgae, k drastically increased with increasing temperature. However, the direct effect of high temperatures on k are obscured by other factors in field experiments compared with laboratory experiments. We defined "fundamental" and "realized" temperature response to explain this effect. Based on relationships for realized temperature response, we predict that proportions of decomposed litter will increase by 0.9%-5% and 4.7%-28.8% by 2100 under low- (2°C) and high-warming conditions (4°C) compared to 2020, respectively. Net litter carbon sinks in BCEs will increase due to higher increase in litter C production than in decomposition by 2100 compared to 2020 under RCP 8.5. We highlight that BCEs will play an increasingly important role in future climate change-mitigation. Our findings can be leveraged for blue carbon accounting under future climate change scenarios.

Modelling blue carbon farming opportunities at different spatial scales.

There is a growing interest in including blue carbon ecosystems (i.e., mangroves, tidal marshes and seagrasses) in climate mitigation programs in national and sub-national policies, with restoration and conservation of these ecosystems identified as potential activities to increase carbon accumulation through time. However, there is still a gap on the spatial scales needed to produce carbon offsets comparable with terrestrial or agricultural ecosystems. Here, we used the Coastal Blue Carbon InVEST 3.7.0 model to estimate future net carbon sequestration in blue carbon ecosystems along Australia's Great Barrier Reef (hereafter GBR) catchments, considering different management scenarios (i.e., reintroduction of tidal exchange through the removal of barriers, sea level rise, restoring low lying land) at three different spatial scales: whole GBR coastline, regional (14,000-16,300 ha), and local (335-370 ha) scales. The focus of the restoration (i.e., tidal marshes and/or mangroves) was dependent on data availability for each scenario. Furthermore, we also estimated the monetary value of carbon sequestration under each management scenario and spatial scale assessed in the study. We found that large scale restoration of tidal marshes could potentially sequester an additional ∼800,000 tonnes of CO2e by 2045 (potentially generating AU$12 million based on the average Australia carbon price), with greater opportunities when sea level rise is accounted for in the modelling. Also, we found that regional and local projects would generate up to 23 tonnes CO2e ha-1 by the end of the crediting period. Our results can guide future decisions in the blue carbon market and financing schemes, however, the return on investment is dependent on the carbon price and funding scheme available for project implementation.

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.

Wave effects on seedling establishment of three pioneer marsh species: survival, morphology and biomechanics.

BACKGROUND AND AIMS: It is important to have an in-depth mechanistic understanding of tidal marsh establishment and dynamics to ensure the long-term persistence of these valuable ecosystems. As wave forcing may be expected to impact seedling establishment, we studied the effect of water-imposed drag forces on seedling survival, morphology and biomechanical properties of three marsh pioneer species that are dominant along the salinity gradient in many areas around the world: Spartina anglica (salt to brackish), Scirpus maritimus (brackish) and Phragmites australis (brackish to fresh). METHODS: Using a newly developed plant-shaking mesocosm (PSM) that mimicked water-imposed wave drag forces, the effect of wave stress on seedling survival was examined, together with impacts on morphology and biomechanical properties. KEY RESULTS: After 7 weeks of exposure to wave stress, lowered seedling survival and growth for all species was revealed. Wave treatments increased the root/shoot biomass ratio to enhance anchorage and made seedlings more flexible (i.e. reduced flexural rigidity), which might be regarded as a mixed outcome between a stress avoidance and stress tolerance strategy. CONCLUSIONS: The different biomechanical responses between the three dominant marsh pioneer species, overall, make them less resistant to external stress. Therefore, our results indicate that the likelihood of marshes becoming established is reduced if wave energy increases. Despite the different biomechanical response of these three pioneer species to waves, the seedlings of all species were found to have low resistance to external stresses.

Long-term monitoring of mercury in adult saltmarsh sparrows breeding in Maine, Massachusetts and New York, USA 2000-2017.

Here we report on the results of a long-term study of mercury exposure in a songbird species, the saltmarsh sparrow (Ammodramus caudacutus). We measured total mercury concentrations in blood (n = 840) and feathers (n = 560) of adult saltmarsh sparrows at six locations between 2000 and 2017: Rachel Carson National Wildlife Refuge (RCNWR) in Wells, Maine; Scarborough Marsh State Wildlife Management Area in Scarborough, Maine; Parker River National Wildlife Refuge on Plum Island, Massachusetts; Pine Neck Preserve in Southampton, Long Island, New York; and North Cinder and North Green Sedge Islands off the coast of Long Island, New York. During the 12-17 year sampling periods, we found that mercury exposure differed by site and year but there was no consistent temporal trend across sites. Blood mercury concentrations declined only at RCNWR in Maine. We also found seasonal variation in blood mercury concentrations and a positive relationship between mercury concentrations of blood and innermost primary feather, but not between blood and tail feather.

Variance reflects resilience to disturbance along a stress gradient: Experimental evidence from coastal marshes.

Quantifying ecosystem resilience to disturbance is important for understanding the effects of disturbances on ecosystems, especially in an era of rapid global change. However, there are few studies that have used standardized experimental disturbances to compare resilience patterns across abiotic gradients in real-world ecosystems. Theoretical studies have suggested that increased return times are associated with increasing variance during recovery from disturbance. However, this notion has rarely been explicitly tested in field, in part due to the challenges involved in obtaining long-term experimental data. In this study, we examined resilience to disturbance of 12 coastal marsh sites (five low-salinity and seven polyhaline [=salt] marshes) along a salinity gradient in Georgia, USA. We found that recovery times after experimental disturbance ranged from 7 to >127 months, and differed among response variables (vegetation height, cover and composition). Recovery rates decreased along the stress gradient of increasing salinity, presumably due to stress reducing plant vigor, but only when low-salinity and polyhaline sites were analyzed separately, indicating a strong role for traits of dominant plant species. The coefficient of variation of vegetation cover and height in control plots did not vary with salinity. In disturbed plots, however, the coefficient of variation (CV) was consistently elevated during the recovery period and increased with salinity. Moreover, higher CV values during recovery were correlated with slower recovery rates. Our results deepen our understanding of resilience to disturbance in natural ecosystems, and point to novel ways that variance can be used either to infer recent disturbance, or, if measured in areas with a known disturbance history, to predict recovery patterns.

Population history and the selective landscape shape patterns of osmoregulatory trait divergence in tidal marsh Savannah sparrows (Passerculus sandwichensis).

A persistent challenge in making associations between phenotypic and environmental variation is understanding how ecological factors and demographic history interact to shape adaptive outcomes. Evaluating the degree to which conspecific populations exposed to similar environmental pressures respond in parallel provides a powerful framework for addressing this challenge. We took this comparative approach with multiple populations of Savannah sparrows (Passerculus sandwichensis) found in tidal marshes along the Pacific coast of North America. The high salinities characterizing tidal marshes select for increased osmoregulatory performance and salinity tolerance. We collected data on physiological traits associated with osmoregulatory performance from 10 tidal marsh and three freshwater-adapted interior populations to evaluate the degree of parallel divergence across populations. All traits showed differences in the magnitude of divergence, but only total evaporative water loss (TEWL) showed differences in the direction of divergence. The drivers of these differences in both the magnitude and direction of divergence varied among traits. For kidney morphology and TEWL, patterns of divergence were best explained by variation in immigration rate from interior populations. Maximum temperature was the best predictor of variation in urine excretion ability, and both gene flow and temperature contributed to variation in plasma osmolality. Finally, analysis of multitrait divergence patterns indicated that differences in the direction of divergence were best explained by population genetic structure, whereas differences in the magnitude of divergence were explained by environmental differences. Together these results show that the influences of demography and the selective landscape can manifest themselves differently across functionally integrated traits.

Divergent selection and drift shape the genomes of two avian sister species spanning a saline-freshwater ecotone.

The role of species divergence due to ecologically based divergent selection-or ecological speciation-in generating and maintaining biodiversity is a central question in evolutionary biology. Comparison of the genomes of phylogenetically related taxa spanning a selective habitat gradient enables discovery of divergent signatures of selection and thereby provides valuable insight into the role of divergent ecological selection in speciation. Tidal marsh ecosystems provide tractable opportunities for studying organisms' adaptations to selective pressures that underlie ecological divergence. Sharp environmental gradients across the saline-freshwater ecotone within tidal marshes present extreme adaptive challenges to terrestrial vertebrates. Here, we sequence 20 whole genomes of two avian sister species endemic to tidal marshes-the saltmarsh sparrow (Ammospiza caudacutus) and Nelson's sparrow (A. nelsoni)-to evaluate the influence of selective and demographic processes in shaping genome-wide patterns of divergence. Genome-wide divergence between these two recently diverged sister species was notably high (genome-wide F ST = 0.32). Against a background of high genome-wide divergence, regions of elevated divergence were widespread throughout the genome, as opposed to focused within islands of differentiation. These patterns may be the result of genetic drift resulting from past tidal march colonization events in conjunction with divergent selection to different environments. We identified several candidate genes that exhibited elevated divergence between saltmarsh and Nelson's sparrows, including genes linked to osmotic regulation, circadian rhythm, and plumage melanism-all putative candidates linked to adaptation to tidal marsh environments. These findings provide new insights into the roles of divergent selection and genetic drift in generating and maintaining biodiversity.

The Importance of Marine Predators in the Provisioning of Ecosystem Services by Coastal Plant Communities.

Food web theory predicts that current global declines in marine predators could generate unwanted consequences for many marine ecosystems. In coastal plant communities (kelp, seagrass, mangroves, and salt marsh), several studies have documented the far-reaching effects of changing predator populations. Across coastal ecosystems, the loss of marine predators appears to negatively affect coastal plant communities and the ecosystem services they provide. Here, we discuss some of the documented and suspected effects of predators on coastal protection, carbon sequestration, and the stability and resilience of coastal plant communities. In addition, we present a meta-analysis to assess the strength and direction of trophic cascades in kelp forests, seagrasses, salt marshes, and mangroves. We demonstrate that the strength and direction of trophic cascades varied across ecosystem types, with predators having a large positive effect on plants in salt marshes, a moderate positive effect on plants in kelp and mangroves, and no effect on plants in seagrasses. Our analysis also identified that there is a paucity of literature on trophic cascades for all four coastal plant systems, but especially seagrass and mangroves. Our results demonstrate the crucial role of predators in maintaining coastal ecosystem services, but also highlights the need for further research before large-scale generalizations about the prevalence, direction, and strength of trophic cascade in coastal plant communities can be made.

Effects of the addition of nitrogen and sulfate on CH4 and CO2 emissions, soil, and pore water chemistry in a high marsh of the Min River estuary in southeastern China.

Exogenous nitrogen (N) and sulfate (SO42-), resulting from human activity, can strongly influence the emission of CH4 and CO2 from soil ecosystems. Studies have reported the effects of N and SO42- on CH4 and CO2 emissions from inland peatlands and paddies. However, very few studies have presented year-round data on the effects of the addition of N and SO42- on CH4 and CO2 emissions in estuarine marshes. The effects of the addition of N and SO42- on the emission of CH4 and CO2 were investigated in a Cyperus malaccensis marsh in the high tidal flat of the Min River estuary of southeastern China from September 2014 to August 2015. Dissolved NH4Cl, KNO3, and K2SO4 were applied every month, in doses of 24gN/SO42-m-2·yr-1. The emission of CH4 and CO2 showed distinct monthly and seasonal variations. Compared with the control, the addition of NH4Cl and NH4NO3+K2SO4 showed increases in CH4 fluxes (p<0.05), while the effects of the addition of KNO3 and K2SO4 on CH4 were minor (p>0.05). NH4Cl had a positive impact on CO2 emissions (p<0.01), while the addition of KNO3, K2SO4, and NH4NO3+K2SO4 had minor positive impacts, compared to the control (p>0.05). Correlation analysis found that soil sulfate concentration, nitrogen availability and enzyme activity were the dominant factors influencing CH4 and CO2 variation. Our findings suggest that CH4 and CO2 emissions were influenced more by ammonium than by nitrate. We propose that the suppressive effect of additional sulfate on CH4 production is insignificant, due to which the inhibition may be overestimated in the estuarine brackish marsh.