Increasing calcium scarcity along Afrotropical forest succession.

Secondary forests constitute an increasingly important component of tropical forests worldwide. Although cycling of essential nutrients affects recovery trajectories of secondary forests, the effect of nutrient limitation on forest regrowth is poorly constrained. Here we use three lines of evidence from secondary forest succession sequences in central Africa to identify potential nutrient limitation in regrowing forests. First, we show that atmospheric phosphorus supply exceeds demand along forest succession, whereas forests rely on soil stocks to meet their base cation demands. Second, soil nutrient metrics indicate that available phosphorus increases along the succession, whereas available cations decrease. Finally, fine root, foliar and litter stoichiometry show that tissue calcium concentrations decline relative to those of nitrogen and phosphorus during succession. Taken together, these observations suggest that calcium becomes an increasingly scarce resource in central African forests during secondary succession. Furthermore, ecosystem calcium storage shifts from soil to woody biomass over succession, making it a vulnerable nutrient in the wake of land-use change scenarios that involve woody biomass export. Our results thus call for a broadened focus on elements other than nitrogen and phosphorus regarding tropical forest biogeochemical cycles and identify calcium as a scarce and potentially limiting nutrient in an increasingly disturbed and dynamic tropical forest landscape.

Low N2O and variable CH4 fluxes from tropical forest soils of the Congo Basin.

Globally, tropical forests are assumed to be an important source of atmospheric nitrous oxide (N2O) and sink for methane (CH4). Yet, although the Congo Basin comprises the second largest tropical forest and is considered the most pristine large basin left on Earth, in situ N2O and CH4 flux measurements are scarce. Here, we provide multi-year data derived from on-ground soil flux (n = 1558) and riverine dissolved gas concentration (n = 332) measurements spanning montane, swamp, and lowland forests. Each forest type core monitoring site was sampled at least for one hydrological year between 2016 - 2020 at a frequency of 7-14 days. We estimate a terrestrial CH4 uptake (in kg CH4-C ha-1 yr-1) for montane (-4.28) and lowland forests (-3.52) and a massive CH4 release from swamp forests (non-inundated 2.68; inundated 341). All investigated forest types were a N2O source (except for inundated swamp forest) with 0.93, 1.56, 3.5, and -0.19 kg N2O-N ha-1 yr-1 for montane, lowland, non-inundated swamp, and inundated swamp forests, respectively.

Fire-derived phosphorus fertilization of African tropical forests.

Central African tropical forests face increasing anthropogenic pressures, particularly in the form of deforestation and land-use conversion to agriculture. The long-term effects of this transformation of pristine forests to fallow-based agroecosystems and secondary forests on biogeochemical cycles that drive forest functioning are poorly understood. Here, we show that biomass burning on the African continent results in high phosphorus (P) deposition on an equatorial forest via fire-derived atmospheric emissions. Furthermore, we show that deposition loads increase with forest regrowth age, likely due to increasing canopy complexity, ranging from 0.4 kg P ha-1 yr-1 on agricultural fields to 3.1 kg P ha-1 yr-1 on old secondary forests. In forest systems, canopy wash-off of dry P deposition increases with rainfall amount, highlighting how tropical forest canopies act as dynamic reservoirs for enhanced addition of this essential plant nutrient. Overall, the observed P deposition load at the study site is substantial and demonstrates the importance of canopy trapping as a pathway for nutrient input into forest ecosystems.

In-depth analysis of N2O fluxes in tropical forest soils of the Congo Basin combining isotope and functional gene analysis.

Primary tropical forests generally exhibit large gaseous nitrogen (N) losses, occurring as nitric oxide (NO), nitrous oxide (N2O) or elemental nitrogen (N2). The release of N2O is of particular concern due to its high global warming potential and destruction of stratospheric ozone. Tropical forest soils are predicted to be among the largest natural sources of N2O; however, despite being the world's second-largest rainforest, measurements of gaseous N-losses from forest soils of the Congo Basin are scarce. In addition, long-term studies investigating N2O fluxes from different forest ecosystem types (lowland and montane forests) are scarce. In this study we show that fluxes measured in the Congo Basin were lower than fluxes measured in the Neotropics, and in the tropical forests of Australia and South East Asia. In addition, we show that despite different climatic conditions, average annual N2O fluxes in the Congo Basin's lowland forests (0.97 ± 0.53 kg N ha-1 year-1) were comparable to those in its montane forest (0.88 ± 0.97 kg N ha-1 year-1). Measurements of soil pore air N2O isotope data at multiple depths suggests that a microbial reduction of N2O to N2 within the soil may account for the observed low surface N2O fluxes and low soil pore N2O concentrations. The potential for microbial reduction is corroborated by a significant abundance and expression of the gene nosZ in soil samples from both study sites. Although isotopic and functional gene analyses indicate an enzymatic potential for complete denitrification, combined gaseous N-losses (N2O, N2) are unlikely to account for the missing N-sink in these forests. Other N-losses such as NO, N2 via Feammox or hydrological particulate organic nitrogen export could play an important role in soils of the Congo Basin and should be the focus of future research.

Century-long apparent decrease in intrinsic water-use efficiency with no evidence of progressive nutrient limitation in African tropical forests.

Forests exhibit leaf- and ecosystem-level responses to environmental changes. Specifically, rising carbon dioxide (CO2 ) levels over the past century are expected to have increased the intrinsic water-use efficiency (iWUE) of tropical trees while the ecosystem is gradually pushed into progressive nutrient limitation. Due to the long-term character of these changes, however, observational datasets to validate both paradigms are limited in space and time. In this study, we used a unique herbarium record to go back nearly a century and show that despite the rise in CO2 concentrations, iWUE has decreased in central African tropical trees in the Congo Basin. Although we find evidence that points to leaf-level adaptation to increasing CO2 -that is, increasing photosynthesis-related nutrients and decreasing maximum stomatal conductance, a decrease in leaf δ13 C clearly indicates a decreasing iWUE over time. Additionally, the stoichiometric carbon to nitrogen and nitrogen to phosphorus ratios in the leaves show no sign of progressive nutrient limitation as they have remained constant since 1938, which suggests that nutrients have not increasingly limited productivity in this biome. Altogether, the data suggest that other environmental factors, such as increasing temperature, might have negatively affected net photosynthesis and consequently downregulated the iWUE. Results from this study reveal that the second largest tropical forest on Earth has responded differently to recent environmental changes than expected, highlighting the need for further on-ground monitoring in the Congo Basin.

Land-use controls on carbon biogeochemistry in lowland streams of the Congo Basin.

The flux and composition of carbon (C) from land to rivers represents a critical component of the global C cycle as well as a powerful integrator of landscape-level processes. In the Congo Basin, an expansive network of streams and rivers transport and cycle terrigenous C sourced from the largest swathe of pristine tropical forest on Earth. Increasing rates of deforestation and conversion to agriculture in the Basin are altering the current regime of terrestrial-to-aquatic biogeochemical cycling of C. To investigate the role of deforestation on dissolved organic and inorganic C (DOC and DIC, respectively) biogeochemistry in the Congo Basin, six lowland streams that drain catchments of varying forest proportion (12%-77%) were sampled monthly for 1 year. Annual mean concentrations of DOC exhibited an asymptotic response to forest loss, while DIC concentrations increased continuously with forest loss. The isotopic signature of DIC became significantly more enriched with deforestation, indicating a shift in source and processes controlling DIC production. The composition of dissolved organic matter (DOM), as revealed by ultra-high-resolution mass spectrometry, indicated that deforested catchments export relatively more aliphatic and heteroatomic DOM sourced from microbial biomass in soils. The DOM compositional results imply that DOM from the deforested sites is more biolabile than DOM from the forest, consistent with the corresponding elevated stream CO2 concentrations. In short, forest loss results in significant and comprehensive shifts in the C biogeochemistry of the associated streams. It is apparent that land-use conversion has the potential to dramatically affect the C cycle in the Congo Basin by reducing the downstream flux of stable, vascular-plant derived DOC while increasing the transfer of biolabile soil C to the atmosphere.

Mobilization of aged and biolabile soil carbon by tropical deforestation.

In the mostly pristine Congo Basin, agricultural land-use change has intensified in recent years. One potential and understudied consequence of this deforestation and conversion to agriculture is the mobilization and loss of organic matter from soils to rivers as dissolved organic matter. Here, we quantify and characterize dissolved organic matter sampled from 19 catchments of varying deforestation extent near Lake Kivu over a two-week period during the wet season. Dissolved organic carbon from deforested, agriculturally-dominated catchments was older (14C age: ~1.5kyr) and more biolabile than from pristine forest catchments. Ultrahigh-resolution mass spectrometry revealed that this aged organic matter from deforested catchments was energy-rich and chemodiverse, with higher proportions of nitrogen- and sulfur-containing formulae. Given the molecular composition and biolability, we suggest that organic matter from deforested landscapes is preferentially respired upon disturbance, resulting in elevated in-stream concentrations of carbon dioxide. We estimate that while deforestation reduces the overall flux of dissolved organic carbon by ~56%, it does not significantly change the yield of biolabile dissolved organic carbon. Ultimately, the exposure of deeper soil horizons through deforestation and agricultural expansion releases old, previously stable, and biolabile soil organic carbon into the modern carbon cycle via the aquatic pathway.

Increasing Alkalinity Export from Large Russian Arctic Rivers.

Riverine carbonate alkalinity (HCO3- and CO32-) sourced from chemical weathering represents a significant sink for atmospheric CO2. Alkalinity flux from Arctic rivers is partly determined by precipitation, permafrost extent, groundwater flow paths, and surface vegetation, all of which are changing under a warming climate. Here we show that over the past three and half decades, the export of alkalinity from the Yenisei and Ob' Rivers increased from 225 to 642 Geq yr-1 (+185%) and from 201 to 470 Geq yr-1 (+134%); an average rate of 11.90 and 7.28 Geq yr-1, respectively. These increases may have resulted from a suite of changes related to climate change and anthropogenic activity, including higher temperatures, increased precipitation, permafrost thaw, changes to hydrologic flow paths, shifts in vegetation, and decreased acid deposition. Regardless of the direct causes, these trends have broad implications for the rate of carbon sequestration on land and delivery of buffering capacity to freshwater ecosystems and the Arctic Ocean.

High fire-derived nitrogen deposition on central African forests.

Atmospheric nitrogen (N) deposition is an important determinant of N availability for natural ecosystems worldwide. Increased anthropogenic N deposition shifts the stoichiometric equilibrium of ecosystems, with direct and indirect impacts on ecosystem functioning and biogeochemical cycles. Current simulation data suggest that remote tropical forests still receive low atmospheric N deposition due to a lack of proximate industry, low rates of fossil fuel combustion, and absence of intensive agriculture. We present field-based N deposition data for forests of the central Congo Basin, and use ultrahigh-resolution mass spectrometry to characterize the organic N fraction. Additionally, we use satellite data and modeling for atmospheric N source apportionment. Our results indicate that these forests receive 18.2 kg N hectare-1 years-1 as wet deposition, with dry deposition via canopy interception adding considerably to this flux. We also show that roughly half of the N deposition is organic, which is often ignored in N deposition measurements and simulations. The source of atmospheric N is predominantly derived from intensive seasonal burning of biomass on the continent. This high N deposition has important implications for the ecology of the Congo Basin and for global biogeochemical cycles more broadly.

Ancient low-molecular-weight organic acids in permafrost fuel rapid carbon dioxide production upon thaw.

Northern permafrost soils store a vast reservoir of carbon, nearly twice that of the present atmosphere. Current and projected climate warming threatens widespread thaw of these frozen, organic carbon (OC)-rich soils. Upon thaw, mobilized permafrost OC in dissolved and particulate forms can enter streams and rivers, which are important processors of OC and conduits for carbon dioxide (CO2) to the atmosphere. Here, we demonstrate that ancient dissolved organic carbon (DOC) leached from 35,800 y B.P. permafrost soils is rapidly mineralized to CO2. During 200-h experiments in a novel high-temporal-resolution bioreactor, DOC concentration decreased by an average of 53%, fueling a more than sevenfold increase in dissolved inorganic carbon (DIC) concentration. Eighty-seven percent of the DOC loss to microbial uptake was derived from the low-molecular-weight (LMW) organic acids acetate and butyrate. To our knowledge, our study is the first to directly quantify high CO2 production rates from permafrost-derived LMW DOC mineralization. The observed DOC loss rates are among the highest reported for permafrost carbon and demonstrate the potential importance of LMW DOC in driving the rapid metabolism of Pleistocene-age permafrost carbon upon thaw and the outgassing of CO2 to the atmosphere by soils and nearby inland waters.