Freshwater wetland plants respond nonlinearly to inundation over a sustained period.

PREMISE: Wetland plants regularly experience physiological stresses resulting from inundation; however, plant responses to the interacting effects of water level and inundation duration are not fully understood. METHODS: We conducted a mesocosm experiment on two wetland species, sawgrass (Cladium jamaicense) and muhly grass (Muhlenbergia filipes), that co-dominate many freshwater wetlands in the Florida Everglades. We tracked photosynthesis, respiration, and growth at water levels of -10 (control), 10 (shallow), and 35 cm (deep) with reference to soil surface over 6 months. RESULTS: The response of photosynthesis to inundation was nonlinear. Specifically, photosynthetic capacity (Amax ) declined by 25% in sawgrass and by 70% in muhly grass after 1-2 months of inundation. After 4 months, Amax of muhly grass in the deep-water treatment declined to near zero. Inundated sawgrass maintained similar leaf respiration and growth rates as the control, whereas inundated muhly grass suppressed both respiration and growth. At the end of the experiment, sawgrass had similar nonstructural carbohydrate pools in all treatments. By contrast, muhly grass in the deep-water treatment had largely depleted sugar reserves but maintained a similar starch pool as the control, which is critical for post-stress recovery. CONCLUSIONS: Overall, the two species exhibited nonlinear and contrasting patterns of carbon uptake and use under inundation stress, which ultimately defines their strategies of surviving regularly flooded habitats. The results suggest that a future scenario with more intensive inundation, due to the water management and climate change, may weaken the dominance of muhly grass in many freshwater wetlands of the Everglades.

Intensified inundation shifts a freshwater wetland from a CO2 sink to a source.

Climate change has altered global precipitation patterns and has led to greater variation in hydrological conditions. Wetlands are important globally for their soil carbon storage. Given that wetland carbon processes are primarily driven by hydrology, a comprehensive understanding of the effect of inundation is needed. In this study, we evaluated the effect of water level (WL) and inundation duration (ID) on carbon dioxide (CO2 ) fluxes by analysing a 10-year (2008-2017) eddy covariance dataset from a seasonally inundated freshwater marl prairie in the Everglades National Park. Both gross primary production (GPP) and ecosystem respiration (ER) rates showed declines under inundation. While GPP rates decreased almost linearly as WL and ID increased, ER rates were less responsive to WL increase beyond 30 cm and extended inundation periods. The unequal responses between GPP and ER caused a weaker net ecosystem CO2 sink strength as inundation intensity increased. Eventually, the ecosystem tended to become a net CO2 source on a daily basis when either WL exceeded 46 cm or inundation lasted longer than 7 months. Particularly, with an extended period of high-WLs in 2016 (i.e., WL remained >40 cm for >9 months), the ecosystem became a CO2 source, as opposed to being a sink or neutral for CO2 in other years. Furthermore, the extreme inundation in 2016 was followed by a 4-month postinundation period with lower net ecosystem CO2 uptake compared to other years. Given that inundation plays a key role in controlling ecosystem CO2 balance, we suggest that a future with more intensive inundation caused by climate change or water management activities can weaken the CO2 sink strength of the Everglades freshwater marl prairies and similar wetlands globally, creating a positive feedback to climate change.

Water uptake of Alaskan tundra evergreens during the winter-spring transition.

PREMISE OF THE STUDY: The cold season in the Arctic extends over 8 to 9 mo, yet little is known about vascular plant physiology during this period. Evergreen species photosynthesize under the snow, implying that they are exchanging water with the atmosphere. However, liquid water available for plant uptake may be limited at this time. The study objective was to determine whether evergreen plants are actively taking up water while under snow and/or immediately following snowmelt during spring thaw. METHODS: In two in situ experiments, one at the plot level and another at the individual species level, (2)H-labeled water was used as a tracer injected beneath the snow, after which plant stems and leaves were tested for the presence of the label. In separate experiments, excised shoots of evergreen species were exposed to (2)H-labeled water for ∼5 s or 60 min and tested for foliar uptake of the label. KEY RESULTS: In both the plot-level and the species-level experiments, some (2)H-labeled water was found in leaves and stems. Additionally, excised individual plant shoots exposed to labeled water for 60 min took up significantly more (2)H-label than shoots exposed ∼5 s. CONCLUSIONS: Evergreen tundra plants take up water under snow cover, some via roots, but also likely by foliar uptake. The ability to take up water in the subnivean environment allows evergreen tundra plants to take advantage of mild spring conditions under the snow and replenish carbon lost by winter respiration.

Plant phenological responses to a long-term experimental extension of growing season and soil warming in the tussock tundra of Alaska.

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis-idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species-specific and growth form-specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.

A general pattern of trade-offs between ecosystem resistance and resilience to tropical cyclones.

Tropical cyclones drive coastal ecosystem dynamics, and their frequency, intensity, and spatial distribution are predicted to shift with climate change. Patterns of resistance and resilience were synthesized for 4138 ecosystem time series from n = 26 storms occurring between 1985 and 2018 in the Northern Hemisphere to predict how coastal ecosystems will respond to future disturbance regimes. Data were grouped by ecosystems (fresh water, salt water, terrestrial, and wetland) and response categories (biogeochemistry, hydrography, mobile biota, sedentary fauna, and vascular plants). We observed a repeated pattern of trade-offs between resistance and resilience across analyses. These patterns are likely the outcomes of evolutionary adaptation, they conform to disturbance theories, and they indicate that consistent rules may govern ecosystem susceptibility to tropical cyclones.

Ecophysiological analysis of two arctic sedges under reduced root temperatures.

Shoot physiological activity in arctic vascular plants may be controlled by low soil temperatures. While leaves may be exposed to moderate temperatures during the growing season, root temperatures often remain near freezing. In this study, two tundra sedges, Eriophorum vaginatum and Carex bigellowii, were subjected to reduced soil temperatures, and photosynthetic parameters (light saturated photosynthesis A(max), variable to maximal fluorescence and F(v)/F(m) stomatal conductance) and abscisic acid concentrations were determined. Stomatal conductance and A(max) for both E. vaginatum and C. bigellowii strongly decreased with declining soil temperatures. Decreasing soil temperature, however, impacted F(v)/F(m) to a much lesser degree. Root and leaf ABA concentrations increased with decreasing root temperature. These observations support the contention that soil temperature is a significant photosynthetic driving factor in arctic sedges exposed to variable root and shoot temperatures. Because these two species comprise approximately 30% of the vascular ground cover of wet tussock tundra, the soil temperature responses of these sedges potentially scale up to significant effects on ecosystem carbon exchange.

El Niño Southern Oscillation (ENSO) enhances CO2 exchange rates in freshwater Marsh ecosystems in the Florida everglades.

This research examines the relationships between El Niño Southern Oscillation (ENSO), water level, precipitation patterns and carbon dioxide (CO2) exchange rates in the freshwater wetland ecosystems of the Florida Everglades. Data was obtained over a 5-year study period (2009-2013) from two freshwater marsh sites located in Everglades National Park that differ in hydrology. At the short-hydroperiod site (Taylor Slough; TS) and the long-hydroperiod site (Shark River Slough; SRS) fluctuations in precipitation patterns occurred with changes in ENSO phase, suggesting that extreme ENSO phases alter Everglades hydrology which is known to have a substantial influence on ecosystem carbon dynamics. Variations in both ENSO phase and annual net CO2 exchange rates co-occurred with changes in wet and dry season length and intensity. Combined with site-specific seasonality in CO2 exchanges rates, El Niño and La Niña phases magnified season intensity and CO2 exchange rates at both sites. At TS, net CO2 uptake rates were higher in the dry season, whereas SRS had greater rates of carbon sequestration during the wet season. As La Niña phases were concurrent with drought years and extended dry seasons, TS became a greater sink for CO2 on an annual basis (-11 to -110 g CO2 m-2 yr-1) compared to El Niño and neutral years (-5 to -43.5 g CO2 m-2 yr-1). SRS was a small source for CO2 annually (1.81 to 80 g CO2 m-2 yr-1) except in one exceptionally wet year that was associated with an El Niño phase (-16 g CO2 m-2 yr-1). Considering that future climate predictions suggest a higher frequency and intensity in El Niño and La Niña phases, these results indicate that changes in extreme ENSO phases will significantly alter CO2 dynamics in the Florida Everglades.

Diurnal patterns of gas-exchange and metabolic pools in tundra plants during three phases of the arctic growing season.

Arctic tundra plant communities are subject to a short growing season that is the primary period in which carbon is sequestered for growth and survival. This period is often characterized by 24-h photoperiods for several months a year. To compensate for the short growing season tundra plants may extend their carbon uptake capacity on a diurnal basis, but whether this is true remains unknown. Here, we examined in situ diurnal patterns of physiological activity and foliar metabolites during the early, mid, and late growing season in seven arctic species under light-saturated conditions. We found clear diurnal patterns in photosynthesis and respiration, with midday peaks and midnight lulls indicative of circadian regulation. Diurnal patterns in foliar metabolite concentrations were less distinct between the species and across seasons, suggesting that metabolic pools are likely governed by proximate external factors. This understanding of diurnal physiology will also enhance the parameterization of process-based models, which will aid in better predicting future carbon dynamics for the tundra. This becomes even more critical considering the rapid changes that are occurring circumpolarly that are altering plant community structure, function, and ultimately regional and global carbon budgets.

Cyclic occurrence of fire and its role in carbon dynamics along an edaphic moisture gradient in longleaf pine ecosystems.

Fire regulates the structure and function of savanna ecosystems, yet we lack understanding of how cyclic fire affects savanna carbon dynamics. Furthermore, it is largely unknown how predicted changes in climate may impact the interaction between fire and carbon cycling in these ecosystems. This study utilizes a novel combination of prescribed fire, eddy covariance (EC) and statistical techniques to investigate carbon dynamics in frequently burned longleaf pine savannas along a gradient of soil moisture availability (mesic, intermediate and xeric). This research approach allowed us to investigate the complex interactions between carbon exchange and cyclic fire along the ecological amplitude of longleaf pine. Over three years of EC measurement of net ecosystem exchange (NEE) show that the mesic site was a net carbon sink (NEE = -2.48 tonnes C ha(-1)), while intermediate and xeric sites were net carbon sources (NEE = 1.57 and 1.46 tonnes C ha(-1), respectively), but when carbon losses due to fuel consumption were taken into account, all three sites were carbon sources (10.78, 7.95 and 9.69 tonnes C ha(-1) at the mesic, intermediate and xeric sites, respectively). Nonetheless, rates of NEE returned to pre-fire levels 1-2 months following fire. Consumption of leaf area by prescribed fire was associated with reduction in NEE post-fire, and the system quickly recovered its carbon uptake capacity 30-60 days post fire. While losses due to fire affected carbon balances on short time scales (instantaneous to a few months), drought conditions over the final two years of the study were a more important driver of net carbon loss on yearly to multi-year time scales. However, longer-term observations over greater environmental variability and additional fire cycles would help to more precisely examine interactions between fire and climate and make future predictions about carbon dynamics in these systems.

Effects of simulated drought on the carbon balance of Everglades short-hydroperiod marsh.

Hydrology drives the carbon balance of wetlands by controlling the uptake and release of CO2 and CH4 . Longer dry periods in between heavier precipitation events predicted for the Everglades region, may alter the stability of large carbon pools in this wetland's ecosystems. To determine the effects of drought on CO2 fluxes and CH4 emissions, we simulated changes in hydroperiod with three scenarios that differed in the onset rate of drought (gradual, intermediate, and rapid transition into drought) on 18 freshwater wetland monoliths collected from an Everglades short-hydroperiod marsh. Simulated drought, regardless of the onset rate, resulted in higher net CO2 losses net ecosystem exchange (NEE) over the 22-week manipulation. Drought caused extensive vegetation dieback, increased ecosystem respiration (Reco ), and reduced carbon uptake gross ecosystem exchange (GEE). Photosynthetic potential measured by reflective indices (photochemical reflectance index, water index, normalized phaeophytinization index, and the normalized difference vegetation index) indicated that water stress limited GEE and inhibited Reco . As a result of drought-induced dieback, NEE did not offset methane production during periods of inundation. The average ratio of net CH4 to NEE over the study period was 0.06, surpassing the 100-year greenhouse warming compensation point for CH4 (0.04). Drought-induced diebacks of sawgrass (C3 ) led to the establishment of the invasive species torpedograss (C4 ) when water was resupplied. These changes in the structure and function indicate that freshwater marsh ecosystems can become a net source of CO2 and CH4 to the atmosphere, even following an extended drought. Future changes in precipitation patterns and drought occurrence/duration can change the carbon storage capacity of freshwater marshes from sinks to sources of carbon to the atmosphere. Therefore, climate change will impact the carbon storage capacity of freshwater marshes by influencing water availability and the potential for positive feedbacks on radiative forcing.