Disentangling direct and indirect effects of water table drawdown on above‐ and belowground plant litter decomposition: consequences for accumulation of organic matter in boreal peatlands

Pristine peatlands are carbon (C)‐accumulating wetland ecosystems sustained by a high water table (WT) and consequent anoxia that slows down decomposition. Persistent WT drawdown as a response to climate and/or land‐use change affects decomposition either directly through environmental factors such as increased oxygenation, or indirectly through changes in plant community composition. This study attempts to disentangle the direct and indirect effects of WT drawdown by measuring the relative importance of environmental parameters (WT depth, temperature, soil chemistry) and litter type and/or litter chemical quality on the 2‐year decomposition rates of above‐ and belowground litter (altogether 39 litter types). Consequences for organic matter accumulation were estimated based on the annual litter production. The study sites were chosen to form a three‐stage chronosequence from pristine (undrained) to short‐term (years) and long‐term (decades) WT drawdown conditions at three nutrient regimes. The direct effects of WT drawdown were overruled by the indirect effects through changes in litter type composition and production. Short‐term responses to WT drawdown were small. In long‐term, dramatically increased litter inputs resulted in large accumulation of organic matter in spite of increased decomposition rates. Furthermore, the quality of the accumulated matter greatly changed from that accumulated in pristine conditions. Our results show that the shift in vegetation composition as a response to climate and/or land‐use change is the main factor affecting peatland ecosystem C cycle, and thus dynamic vegetation is a necessity in any model applied for estimating responses of C fluxes to changing environment. We provide possible grouping of litter types into plant functional types that the models could utilize. Furthermore, our results clearly show a drop in soil summer temperature as a response to WT drawdown when an initially open peatland converts into a forest ecosystem, which has not yet been considered in the existing models.     

Simple three‐pool model accurately describes patterns of long‐term litter decomposition in diverse climates

As atmospheric CO₂ increases, ecosystem carbon sequestration will largely depend on how global changes in climate will alter the balance between net primary production and decomposition. The response of primary production to climatic change has been examined using well‐validated mechanistic models, but the same is not true for decomposition, a primary source of atmospheric CO₂. We used the Long‐term Intersite Decomposition Experiment Team (LIDET) dataset and model‐selection techniques to choose and parameterize a model that describes global patterns of litter decomposition. Mass loss was best represented by a three‐pool negative exponential model, with a rapidly decomposing labile pool, an intermediate pool representing cellulose, and a recalcitrant pool. The initial litter lignin/nitrogen ratio defined the size of labile and intermediate pools. Lignin content determined the size of the recalcitrant pool. The decomposition rate of all pools was modified by climate, but the intermediate pool's decomposition rate was also controlled by relative amounts of litter cellulose and lignin (indicative of lignin‐encrusted cellulose). The effect of climate on decomposition was best represented by a composite variable that multiplied a water‐stress function by the Lloyd and Taylor variable Q₁₀ temperature function. Although our model explained nearly 70% of the variation in LIDET data, we observed systematic deviations from model predictions. Below‐ and aboveground material decomposed at notably different rates, depending on the decomposition stage. Decomposition in certain ecosystem‐specific environmental conditions was not well represented by our model; this included roots in very wet and cold soils, and aboveground litter in N‐rich and arid sites. Despite these limitations, our model may still be extremely useful for global modeling efforts, because it accurately (R²=0.6804) described general patterns of long‐term global decomposition for a wide array of litter types, using relatively minimal climatic and litter quality data.     

Decoupling the direct and indirect effects of climate on plant litter decomposition: Accounting for stress‐induced modifications in plant chemistry

Decomposition of plant litter is a fundamental ecosystem process that can act as a feedback to climate change by simultaneously influencing both the productivity of ecosystems and the flux of carbon dioxide from the soil. The influence of climate on decomposition from a postsenescence perspective is relatively well known; in particular, climate is known to regulate the rate of litter decomposition via its direct influence on the reaction kinetics and microbial physiology on processes downstream of tissue senescence. Climate can alter plant metabolism during the formative stage of tissues and could shape the final chemical composition of plant litter that is available for decomposition, and thus indirectly influence decomposition; however, these indirect effects are relatively poorly understood. Climatic stress disrupts cellular homeostasis in plants and results in the reprogramming of primary and secondary metabolic pathways, which leads to changes in the quantity, composition, and organization of small molecules and recalcitrant heteropolymers, including lignins, tannins, suberins, and cuticle within the plant tissue matrix. Furthermore, by regulating metabolism during tissue senescence, climate influences the resorption of nutrients from senescing tissues. Thus, the final chemical composition of plant litter that forms the substrate of decomposition is a combined product of presenescence physiological processes through the production and resorption of metabolites. The changes in quantity, composition, and localization of the molecular construct of the litter could enhance or hinder tissue decomposition and soil nutrient cycling by altering the recalcitrance of the lignocellulose matrix, the composition of microbial communities, and the activity of microbial exo‐enzymes via various complexation reactions. Also, the climate‐induced changes in the molecular composition of litter could differentially influence litter decomposition and soil nutrient cycling. Compared with temperate ecosystems, the indirect effects of climate on litter decomposition in the tropics are not well understood, which underscores the need to conduct additional studies in tropical biomes. We also emphasize the need to focus on how climatic stress affects the root chemistry as roots contribute significantly to biogeochemical cycling, and on utilizing more robust analytical approaches to capture the molecular composition of tissue matrix that fuel microbial metabolism.     

Evaluating litter decomposition in earth system models with long‐term litterbag experiments: an example using the Community Land Model version 4 (CLM4)

Decomposition is a large term in the global carbon budget, but models of the earth system that simulate carbon cycle‐climate feedbacks are largely untested with respect to litter decomposition. We tested the litter decomposition parameterization of the community land model version 4 (CLM4), the terrestrial component of the community earth system model, with data from the long‐term intersite decomposition experiment team (LIDET). The LIDET dataset is a 10‐year study of litter decomposition at multiple sites across North America and Central America. We performed 10‐year litter decomposition simulations comparable with LIDET for 9 litter types and 20 sites in tundra, grassland, and boreal, conifer, deciduous, and tropical forest biomes using the LIDET‐provided climatic decomposition index to constrain temperature and moisture effects on decomposition. We performed additional simulations with DAYCENT, a version of the CENTURY model, to ask how well an established ecosystem model matches the observations. The results show large discrepancy between the laboratory microcosm studies used to parameterize the CLM4 litter decomposition and the LIDET field study. Simulated carbon loss is more rapid than the observations across all sites, and nitrogen immobilization is biased high. Closer agreement with the observations requires much lower decomposition rates, obtained with the assumption that soil mineral nitrogen severely limits decomposition. DAYCENT better replicates the observations, for both carbon mass remaining and nitrogen, independent of nitrogen limitation. CLM4 has low soil carbon in global earth system simulations. These results suggest that this bias arises, in part, from too rapid litter decomposition. More broadly, the terrestrial biogeochemistry of earth system models must be critically tested with observations, and the consequences of particular model choices must be documented. Long‐term litter decomposition experiments such as LIDET provide a real‐world process‐oriented benchmark to evaluate models.     

Estimating the contribution of leaf litter decomposition to soil CO2 efflux in a beech forest using 13C-depleted litter

The contribution of leaf litter decomposition to total soil CO₂ efflux (F_L/F) was evaluated in a beech (Fagus sylvatica L.) forest in eastern France. The Keeling‐plot approach was applied to estimate the isotopic composition of respired soil CO₂ from soil covered with either control (?30.32‰) or ¹³C‐depleted leaf litter (?49.96‰). The δ¹³C of respired soil CO₂ ranged from ?25.50‰ to ?22.60‰ and from ?24.95‰ to ?20.77‰, respectively, with depleted or control litter above the soil. The F_L/F ratio was calculated by a single isotope linear mixing model based on mass conservation equations. It showed seasonal variations, increasing from 2.8% in early spring to about 11.4% in mid summer, and decreasing to 4.2% just after leaf fall. Between December 2001 and December 2002, cumulated F and F_L reached 0.98 and 0.08 kg_C m^?2, respectively. On an annual basis, decomposition of fresh leaf litter accounted for 8% of soil respiration and 80% of total C loss from fresh leaf litter. The other fraction of carbon loss during leaf litter decomposition that is assumed to have entered the soil organic matter pool (i.e. 20%) represents only 0.02 kg_C m^?2.     

Annual temperature variation as a time machine to understand the effects of long‐term climate change on a poleward range shift

Range shifts due to annual variation in temperature are more tractable than range shifts linked to decadal to century long temperature changes due to climate change, providing natural experiments to determine the mechanisms responsible for driving long‐term distributional shifts. In this study we couple physiologically grounded mechanistic models with biogeographic surveys in 2 years with high levels of annual temperature variation to disentangle the drivers of a historical range shift driven by climate change. The distribution of the barnacle Semibalanus balanoides has shifted 350 km poleward in the past half century along the east coast of the United States. Recruits were present throughout the historical range following the 2015 reproductive season, when temperatures were similar to those in the past century, and absent following the 2016 reproductive season when temperatures were warmer than they have been since 1870, the earliest date for temperature records. Our dispersal dependent mechanistic models of reproductive success were highly accurate and predicted patterns of reproduction success documented in field surveys throughout the historical range in 2015 and 2016. Our mechanistic models of reproductive success not only predicted recruitment dynamics near the range edge but also predicted interior range fragmentation in a number of years between 1870 and 2016. All recruits monitored within the historical range following the 2015 colonization died before 2016 suggesting juvenile survival was likely the primary driver of the historical range retraction. However, if 2016 is indicative of future temperatures mechanisms of range limitation will shift and reproductive failure will lead to further range retraction in the future. Mechanistic models are necessary for accurately predicting the effects of climate change on ranges of species.     

Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water‐use efficiency at the leaf‐ and ecosystem‐scales in a northern Great Plains grassland

Measurements of the carbon (δ¹³C_m) and oxygen (δ¹⁸O_m) isotope composition of C₃ plant tissue provide important insights into controls on water‐use efficiency. We investigated the causes of seasonal and inter‐annual variability in water‐use efficiency in a grassland near Lethbridge, Canada using stable isotope (leaf‐scale) and eddy covariance measurements (ecosystem‐scale). The positive relationship between δ¹³C_m and δ¹⁸O_m values for samples collected during 1998–2001 indicated that variation in stomatal conductance and water stress‐induced changes in the degree of stomatal limitation of net photosynthesis were the major controls on variation in δ¹³C_m and biomass production during this time. By comparison, the lack of a significant relationship between δ¹³C_m and δ¹⁸O_m values during 2002, 2003 and 2006 demonstrated that water stress was not a significant limitation on photosynthesis and biomass production in these years. Water‐use efficiency was higher in 2000 than 1999, consistent with expectations because of greater stomatal limitation of photosynthesis and lower leaf c_i/ca during the drier conditions of 2000. Calculated values of leaf‐scale water‐use efficiency were 2–3 times higher than ecosystem‐scale water‐use efficiency, a difference that was likely due to carbon lost in root respiration and water lost during soil evaporation that was not accounted for by the stable isotope measurements.