Fine Root Production across a Primary Successional Ecosystem Chronosequence at Mt. Shasta,California

Estimating changes in belowground biomass and production is essential for understanding fundamental patterns and processes during ecosystem development. We examined patterns of fine root production, aboveground litterfall, and forest floor accumulation during forest primary succession at the Mt. Shasta Mudflows ecosystem chronosequence. Fine root production was measured using the root ingrowth cores method over 1 year, and aboveground litterfall was collected over 2 years. Fine root production increased significantly with ecosystem age, but only the youngest ecosystem was significantly different from all of the older ecosystems. Root production was 44.5 ± 13.3, 168.3 ± 20.6, 190.5 ± 33.8, and 236.3 ± 65.4 g m⁻² y⁻¹ in the 77, 255, 616, and >850-year-old ecosystems, respectively. Generally, aboveground litterfall and forest floor accumulation did not follow the same pattern as root production. The relative contribution of fine root production to total fine detrital production increased significantly with ecosystem age, from 14 to 49%, but only the youngest ecosystem was significantly different from all of the older ecosystems. Fine root production was significantly correlated with some measures of soil fertility but was not correlated with leaf or total litterfall, or forest floor accumulation. It was best predicted by soil N concentration alone, but this relationship may not be causal, as soil N concentration was also correlated with ecosystem age. For the oldest ecosystem, fine root production was also measured using the sequential intact cores/compartment-flow model method, and the difference between the two estimates was not significant. Our study suggests that the relative contribution of fine roots to fine detrital production, and hence to soil organic matter accumulation, may increase during forest primary succession.     

Dissolved organic matter under native Cerrado and Pinus caribaea plantations in the Brazilian savanna

The transformation of native Cerrado into Pinus caribaea Morelet plantations changes the DOM dynamics including changed rates of mineralisation, denitrification, and C export to the groundwater. To examine the differences in quantity, temporal dynamics, and quality of DOM between Cerrado and Pine plantations we collected rainfall, throughfall, stemflow, litter leachate (under pine only) and soil solution at 15, 80, and 200 cm depth in weekly intervals during the rainy seasons 1997/98 and 1998/99. We determined total dissolved organic carbon (DOC) concentrations and assessed DOM quality by separating hydrophilic and hydrophobic fractions and by NMR analysis of organic layer extracts. The rainfall had a mean DOC concentration of 2.6 mg L^–1. The mean concentrations of DOC in the throughfall of the pine plantations (5.0–10.5 mg L^–1) were significantly above those of Cerrado (2.6–4.9 mg L^–1). During the first part of the rainy seasons (October–December), the concentrations of DOC in the soil solution (15–200 cm depth) under Cerrado and pine did not differ significantly. During the second part of both rainy seasons (January–April), the concentrations of DOC in the soil solution under Cerrado (4.4–5.1 mg L^–1) exceeded those under PI (1.4–2.7 mg L^–1). Possible explanations of the latter include higher DOM input into the Cerrado soil and a stronger retention and/or faster mineralisation of the pine DOM than of the Cerrado DOM in the mineral soil. As the structural composition of DOM extracted from the organic layer under Cerrado and pine did not differ significantly, faster mineralisation was the most likely explanation for partly lower DOC concentrations in the soil solution under pine than under Cerrado. This assumption was supported by increasing contributions of hydrophobic DOM to total DOM with increasing depth under pine while, under Cerrado, the DOM composition did not change with depth. The reason for DOM mineralisation under pine was probably the higher N availability because total N concentrations were 11–23 times higher under pine than under Cerrado.     

Production of Total Potentially Soluble Organic C, N, and P Across an Ecosystem Chronosequence: Root versus Leaf Litter

Dissolved organic matter (DOM) plays several important roles in forest ecosystem development, undergoing chemical, physical and/or biological reactions that affect ecosystem nutrient retention. Very few studies have focused on gross rates of DOM production, and we know of no study that has directly measured DOM production from root litter. Our objectives were to quantify major sources of total potentially water-soluble organic matter (DOM_tps) production, with an emphasis on production from root litter, to quantify and compare total potentially soluble organic C, N, and P (DOC_tps, DON_tps, and DOP_tps) production, and to quantify changes in their production during forest primary succession and ecosystem development at the Mt. Shasta Mudflows ecosystem chronosequence. To do so, we exhaustively extracted freshly senesced root and leaf and other aboveground litter for DOC_tps, DON_tps, and DOP_tps by vegetation category, and we calculated DOM_tps production (g m⁻² y⁻¹) at the ecosystem level using data for annual production of fine root and aboveground litter. DOM production from throughfall was calculated by measuring throughfall volume and concentration over 2 years. Results showed that DOM_tps production from root litter was a very important source of DOM_tps in the Mount Shasta mudflow ecosystems, in some cases comparable to production from leaf litter for DON_tps and larger than production from leaf litter for DOP_tps. Total DOC_tps and DON_tps production from all sources increased early in succession from the 77- to the 255-year-old ecosystem. However, total DOP_tps production across the ecosystem chronosequence showed a unique pattern. Generally, the relative importance of root litter for total fine detrital DOC_tps and DON_tps production increased significantly during ecosystem development. Furthermore, DOC_tps and DON_tps production were predominantly driven by changes in biomass production during ecosystem development, whereas changes in litter solubility due to changes in species composition had a smaller effect. We suggest that DOM_tps production from root litter may be an important source of organic matter for the accumulation of SOM during forest ecosystem development. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Shauna M. Uselman, Robert G. Qualls, and Juliane Lilienfein conceived of or designed the study and performed research. SMU analyzed data and wrote the article. SMU and RGQ contributed new methods or models.     

Element storage in native, agri-, and silvicultural ecosystems of the Brazilian savanna

The expanding agriculture in the Brazilian savanna, the Cerrado, changes C and nutrient storages of the savanna ecosystems thereby affecting the global C budget and the sustainability of the local land use. We examined the biomass and the C, N, P, and S storages in above- and belowground biomass, in the organic layer, and in the top 2 m of the mineral soil (Anionic Acrustoxes) of three replicate plots of each of native Cerrado, Pinus caribaea Morelet plantations, productive and degraded Bracchiaria decumbens Stapf. pastures, and of conventional and no-tillage soybean cultivation. Aboveground biomass – in the cropping systems shortly before harvest – decreased in the order, Pinus (15 kg m^–2) > Cerrado (2.3) > conventional tillage (1.9) > no tillage (1.5) > productive pasture (0.64) > degraded pasture (0.37) and belowground biomass in the order, Pinus (9.1) > Cerrado (3.0) > productive pasture (2.2) > degraded pasture (1.5) > conventional tillage (0.60) > no tillage (0.41). The aboveground biomass contained 1.1 (degraded pasture) to 19% (Pinus) of the total C storage, 0.3 (productive pasture, degraded pasture) to 3.5% of the total N storage, 0.3 (degraded pasture) to 2.1% (no tillage, conventional tillage) of the total P storage, and 0.3 (degraded pasture) to 3.7% (Pinus) of the total S storage of the ecosystems. Total C storage in the ecosystems was significantly larger in the Pinus stands (36 kg m^–2) than in all other systems; differences among Cerrado (20), degraded pasture (19), productive pasture (20), no tillage (19), and conventional tillage (19) were small and not significant. All land-use systems had larger N (Pinus, 1.5; degraded pasture, 1.3; productive pasture, 1.4; no tillage, 1.4; conventional tillage, 1.4 kg m^–2) and S storage (PI, 28; degraded pasture, 33; productive pasture, 34; no tillage, 36; conventional tillage, 38 g m^–2) than the Cerrado (N, 1.2 kg; S, 26 g m^–2). The P storages varied between 17 and 29 g m^–2 and were not significantly different among the studied ecosystems. The N and S accumulations in the 12–20-year-old land-use systems were larger than the cumulative known fertilizer inputs indicating that there were unknown inputs possibly including the exploration of the deeper subsoil by deep-reaching roots and transfer of nutrients to the topsoil. Our results indicate that afforestation with Pinus trees has the potential to sequester large amounts of C while pasture degradation, no tillage, and conventional tillage tended to result in small C losses. Land use resulted in a marked accumulation of N and S relative to the Cerrado.     

Biogeochemical consequences of the transformation of native Cerrado into Pinus caribaea plantations in Brazil

Under the same climatic and edaphic conditions, native savanna vegetation in Brazil, the Cerrado, shows a lower stature and canopy cover than planted Pinus caribaea Morelet forests. To assess the differences in biogeochemical element cycling we compared the nutrient economy of Cerrado and Pinus on three replicate plots of each forest type. The mean nutrient storage in the soil organic layer under Pinus (N: 2630; P: 141; K: 103; Ca: 131; Mg: 20 kg ha^–1) was substantially higher than under Cerrado (N: 23; P: 1.2; K: 0.83; Ca: 5.8; Mg: 1.0 kg ha^–1) probably because the Pinus roots explored a larger soil volume. The Pinus trees had a higher nutrient-use efficiency as indicated by higher mean litter mass per unit nutrient in litter (N: 108; P: 2290; K: 729; Ca: 1360; Mg: 5420; S: 1190; Fe: 2960; Mn: 9990, Zn: 145000) than the Cerrado trees (N: 94; P: 1810; K: 619; Ca: 302; Mg: 938, S: 746; Fe: 1800; Mn: 7880; Zn: 63700). Mean annual small litterfall collected in 0.25-m² samplers between May 1997 and April 1999 was 2.1 Mg ha^–1 in Cerrado and 7.8 in Pinus. The litterfall rates of the 1–3 week collection intervals correlated negatively with the soil matric potential indicating that litterfall was partly related to water stress. The fluxes of N (73 kg ha^–1 year^–1), P (3.7), K (11), S (7.0), and Mn (0.83) to the soil with litterfall under Pinus were greater than the litterfall+turnover of the grass/herbs layer under Cerrado (N: 39, P: 2.8, K: 8.6, S: 5.4, Mn: 0.79 kg ha^–1 year^–1), those of Zn (0.06–0.07) were similar, and those of Ca (Pinus: 5.9/Cerrado: 10), Mg (1.5/4.4), and Fe (2.9/4.0) were smaller. Mean residence times of the organic matter and of all elements were longer in the soil organic layer under Pinus (3.7–26 years in the Oi horizon, 8.1–907 years in the whole organic layer) than under Cerrado (0.22–3.6 years in the Oi horizon, the only organic horizon under Cerrado). Our results demonstrate that the main differences in biogeochemical element cycling between the Pinus forest and the Cerrado consisted of a larger nutrient storage in the organic layer, a higher nutrient-use efficiency, and slower nutrient release rates from the organic layer in the Pinus forest than in the Cerrado. Nutrient cycling as assessed by the nutrient fluxes with litterfall was only partly faster in the Pinus forest than in the Cerrado.     

Element storage in native,agri-, and silvicultural ecosystems of the Brazilian savanna. II. Metals

Conversion of native savanna in Brazil, the Cerrado, to agri- and silvicultural land use causes changes in metal storages of the ecosystems. To evaluate the sustainability of land use these changes have to be known. Therefore, we examined the Al, Ca, Fe, K, Mg, Mn, Na, and Zn storages in above- and belowground biomass, the organic layer, and the top 2 m of the mineral soil (Anionic Acrustoxes) of three replicate plots in each of six native and land-use systems. The systems were native Cerrado, Pinus caribaea Morelet plantations, productive and degraded Brachiaria decumbens Stapf pastures, and conventional and no-tillage soybean cultivation. The total metal storage varied little among the studied systems except for Ca, K, and Mg. All land-use systems had larger Ca storages (cropping systems 202–205 g m^–2, productive pasture: 112, degraded pasture: 84, Pinus: 81) than the Cerrado (62 g m^–2). The K storage was smaller in the pastures (17–18 g m^–2) than in Cerrado and Pinus stands (22–24) and largest in the cropping systems (26). The Mg storages were largest in the cropping systems (65–69) and productive pasture (59 g m^–2); those in the Pinus stands (52), the degraded pasture (51), and the Cerrado (53) were similar. For most metals, the aboveground biomass contained up to 1% of the total storage including the top 2 m of the soil (<5% if the lower ecosystem boundary was set at 0.3 m soil depth). However, the aboveground biomass stored up to 12% of Ca, K, and Mg down to 2 m soil depth (41% if the lower ecosystem boundary was set at 0.3 m soil depth). In the Pinus stands, the storage of most metals was larger in the below- than in the aboveground biomass; for the other systems the reverse was true. Metal storages in soil were little affected by land use except that liming resulted in increased Ca and Mg storages in the topsoil. The comparison between known inputs of Ca, K, and Mg and mean annual change rates of their storages revealed that there were considerable base metal losses by leaching, grazing, and removal with the harvest. After 12–20 years, the land-use impact on metal storages is restricted to Ca, Mg, and K. Generally, all land-use systems tend to be richer in these nutrients except for the significant depletion in K of the pastures.     

Water and element input into native, agri- and silvicultural ecosystems of the Brazilian savanna

The interaction of rain water with the vegetation canopy results in changes of the water quantity and quality. We examined these canopy effects in different ecosystems of the Brazilian savanna, the Cerrado. The ecosystems were 20 yr-old Pinus caribaea Morelet plantations (PI), productive (PP) and degraded Brachiaria decumbens Stapf pastures (DP), continuous corn-soybean rotation (CC), and native typical cerrado (CE). We collected rainfall, throughfall, and, in PI and CE, stemflow from three plots of each ecosystem. Dry deposition and canopy leaching were estimated with a Na-tracer method. Between May 1997 and April 1999, the mean annual rainfall was 1656 mm of which 145 mm fell during the dry season (May–September). The throughfall percentage of the rainfall increased in the order, PI (75–85%) < CC (76–89%) < CE (89–100%) < PP (90–100%) < DP (99–100%); stemflow was < 1% of the rainfall. The volume-weighted mean (VWM) pH in rainfall was higher in the dry (6.5) than in the rainy season (5.4). The VWM pH in throughfall decreased in the order, CC (rainy season: 5.9/dry season: 6.2) > PP (5.5/6.0) > CE (5.2/6.0) > DP (5.2/5.6) > PI (4.8/5.7). The rainfall deposition of the dry season contributed one third of the annual element input with rainfall because of higher element concentrations than in the rainy season. The mean Na deposition ratios, i.e. the ratio of throughfall (+ stemflow) to rainfall deposition as a measure for dry deposition, increased in the order, CE (1.5) = CC (1.5) < PP (1.7) < PI (1.9) < (DP 2.1). Total deposition (rainfall + dry deposition) accounted for 104–164% of the K and Ca fertilizer application in PP and for 6.1–12% of the K, Ca, and Mg fertilizer application in CC. The P concentrations were below the detection limit of 0.2 mg L^–1 in all samples. Net canopy uptake, i.e. a smaller throughfall(+ stemflow) than rainfall + dry deposition, of Ca, K, Mg, S, Cu, and Zn in at least one of CE, PI, DP, and PP indicate that plant growth may be limited in part by these nutrients. During the vegetation period, between 28 and 50% of the applied K and Ca were leached from the canopy in PP and between 8.7 and 17% of the applied K, Ca, Mg, and S in CC. Our results demonstrate that PI causes larger water losses and enhanced acid inputs to the soil compared with all other ecosystems. However, the PI and pasture canopies scavenge more nutrients from the atmosphere than CE and CC.