Pine Forest Floor Carbon Accumulation in Response to N and PK Additions: Bomb <Superscript>14</Superscript>C Modelling and Respiration Studies

The addition of nitrogen via deposition alters the carbon balance of temperate forest ecosystems by affecting both production and decomposition rates. The effects of 20 years of nitrogen (N) and phosphorus and potassium (PK) additions were studied in a 40-year-old pine stand in northern Sweden. Carbon fluxes of the forest floor were reconstructed using a combination of data on soil ¹⁴C, tree growth, and litter decomposition. N-only additions caused an increase in needle litterfall, whereas both N and PK additions reduced long-term decomposition rates. Soil respiration measurements showed a 40% reduction in soil respiration for treated compared to control plots. The average age of forest floor carbon was 17 years. Predictions of future soil carbon storage indicate an increase of around 100% in the next 100 years for the N plots and 200% for the NPK plots. As much as 70% of the increase in soil carbon was attributed to the decreased decomposition rate, whereas only 20% was attributable to increased litter production. A reduction in decomposition was observed at a rate of N addition of 30 kg C ha^–1 y^–1, which is not an uncommon rate of N deposition in central Europe. A model based on the continuous-quality decomposition theory was applied to interpret decomposer and substrate parameters. The most likely explanations for the decreased decomposition rate were a fertilizer-induced increase in decomposer efficiency (production-to-assimilation ratio), a more rapid rate of decrease in litter quality, and a decrease in decomposer basic growth rate.     

Effects of experimental fuel additions on fire intensity and severity: unexpected carbon resilience of a neotropical forest

Global changes and associated droughts, heat waves, logging activities, and forest fragmentation may intensify fires in Amazonia by altering forest microclimate and fuel dynamics. To isolate the effects of fuel loads on fire behavior and fire‐induced changes in forest carbon cycling, we manipulated fine fuel loads in a fire experiment located in southeast Amazonia. We predicted that a 50% increase in fine fuel loads would disproportionally increase fire intensity and severity (i.e., tree mortality and losses in carbon stocks) due to multiplicative effects of fine fuel loads on the rate of fire spread, fuel consumption, and burned area. The experiment followed a fully replicated randomized block design (N = 6) comprised of unburned control plots and burned plots that were treated with and without fine fuel additions. The fuel addition treatment significantly increased burned area (+22%) and consequently canopy openness (+10%), fine fuel combustion (+5%), and mortality of individuals ≥5 cm in diameter at breast height (dbh; +37%). Surprisingly, we observed nonsignificant effects of the fuel addition treatment on fireline intensity, and no significant differences among the three treatments for (i) mortality of large trees (≥30 cm dbh), (ii) aboveground forest carbon stocks, and (iii) soil respiration. It was also surprising that postfire tree growth and wood increment were higher in the burned plots treated with fuels than in the unburned control. These results suggest that (i) fine fuel load accumulation increases the likelihood of larger understory fires and (ii) single, low‐intensity fires weakly influence carbon cycling of this primary neotropical forest, although delayed postfire mortality of large trees may lower carbon stocks over the long term. Overall, our findings indicate that increased fine fuel loads alone are unlikely to create threshold conditions for high‐intensity, catastrophic fires during nondrought years.     

Long-term soil nutrient dynamics and lateral nutrient movement in fertilized and unfertilized red pine plantations

In this study, we use arepeated-measures analysis to test thehypothesis that soil fertility underpotassium-limited red pine (Pinusresinosa Ait.) stands at the Charles LathropPack Demonstration Forest in Warrensburg, NewYork is increasing toward a steady state thatwas artificially induced in fertilized standsby K-fertilization over 50 years ago. Wemeasured soil K by horizon and added new datato a 53-year database. We examine onemechanism that explains the higher rate of Kaccumulation in unfertilized stands comparedwith fertilized – lateral movement offertilizer K from treated plots to untreated –using the rubidium/potassium reverse tracermethod. Over the past five decades, soil Kconcentrations under both fertilized andunfertilized red pine have increasedsignificantly. The trends under fertilized andunfertilized plots demonstrate the gradualconvergence of soil K under unfertilized plotstoward concentrations in fertilized plots. Five decades after fertilization, treated soilsstill contain greater concentrations ofexchangeable K and lower bulk densities thanunfertilized plots. Analysis of Rb/K ratios inthe forest floor of fertilized and unfertilizedplots confirms the hypothesis that lateraltransport of surface broadcast fertilizer,applied over 50 years ago, extendsapproximately 11–16 m from the edges offertilized plots. The four unfertilized plotsclosest to fertilized plots have beensignificantly affected by inputs of fertilizerK, while the remaining five plots arerelatively unimpacted. Approximately 36% ofthe K in fertilized plots, and 23% of the K inunfertilized plots affected by fertilizermigration were derived directly from thefertilizer applied 5 decades ago, demonstratingthe highly conservative nature of mineralnutrient cycling in aggrading forests.     

Short-term increases in relative nitrification rates due to trenching in forest floor and mineral soil horizons of different forest types

The representation of NO₃ ^– dynamics within forest growth simulation models could improve forest management. An extensive literature review revealed an 88% probability of measuring a higher relative nitrification index (i.e. RNI = [NO₃ ^–] ÷ [NO₃ ^– + NH₄ ⁺]) in mineral soil horizons than in forest floors, across a wide range of conifer and hardwood ecosystems. We then hypothesised that humus form and fine root density could be used as two crude variables to predict changes in in situ, potential and relative nitrification rates. Twenty-seven trench plots were established in 1999, across nine contrasting hardwood and coniferous stands in the Eastern Townships of Québec. Forest floor and mineral soil samples were collected from each plot, and from a 1 m radius surrounding each plot, on three dates during summer 2000. In situRNI values increased significantly in trench plots as the season progressed. Potential nitrification rates (i.e. NO₃ ^– concentrations following incubation) were two orders of magnitude higher in forest floor than in mineral soil samples. RNI was significantly higher in mineral soil than in forest floor samples after incubations, but the relative increase in RNI due to trenching was higher in forest floor samples. Indices of available C were significantly higher in forest floor than mineral soil samples, and decreased only in forest floor samples during incubations. Likewise, trenching significantly reduced available C in forest floor samples only. Seasonal changes in soil temperature and fine root growth were the most plausible explanations for seasonal changes in NO₃ ^– dynamics, whereas other factors such soil acidity and moisture appeared less important in determining NO₃ ^– dynamics in this study. We conclude that crude assessments of humus form and fine root density have the potential to be used as calibration parameters for the simulation of NO₃ ^– dynamics in forest growth and yield models.     

Effect of typhoon disturbance on fine litterfall and related nutrient input in a subtropical forest on Okinawa Island, Japan

Typhoons are frequent on Okinawa Island, southwestern Japan. The effects of typhoon disturbance on the patterns of fine litterfall and related nutrient inputs in a subtropical evergreen broad-leaved forest were studied over 5 years from May 1996 to April 2001. Annual fine litterfall averaged 7558 kg ha⁻¹ (range from 6188 to 9439 kg ha⁻¹) for six sampling plots over 5 years, which differed significantly among years (p<0.001) but not among plots (p=0.122). A seasonal maximum was most evident for leaf litter component. Woody litter fell more irregularly through the year, and peak fall varied with typhoon and windstorm. The mean ratio of annual litterfall mass of sexual organs to leaves was 0.06, much lower than that in other tropical and subtropical rain forests. Nutrient concentrations varied in litterfall components, but were not significantly different among plots. The lowest concentrations of N and P in leaf litter were observed in March, which is also the month with the greatest leaf fall. However, the highest concentrations were recorded in typhoon season. Nitrogen and P concentrations were 34% and 106% greater in the green leaves that fell during typhoons than in senescent leaves. Mean nutrient inputs by litterfall were: N 83, P 3.2, K 25, Ca 71, Mg 19, Al 12, Na 10, Fe 0.86 and Mn 3.9 kg ha⁻¹ yr⁻¹, and differed significantly among years for all elements (p<0.0005) and among plots only for K (p<0.05) and Mn (p<0.0001). Typhoon disturbance strongly affected annual fine litterfall and related nutrient inputs, which contributed an average of 30% of the annual litterfall mass, and from 30% to 39% (for different nutrient elements) of annual total nutrient inputs. The results from this study suggest that typhoon-driven maintenance of rapid cycling of P and N and their high availability in soil appears to be an important mechanism to maintain productivity in the subtropical forest on Okinawa Island.     

Effects of liming and boron fertilization on boron uptake of Picea abies

The effects of liming on concentrations of boron and other elements in Norway spruce [Picea abies (L) Karst.] needles and in the mor humus layer were studied in long-term field experiments with and without B fertilizer on podzolic soils in Finland. Liming (2000+4000 kg ha^-1 last applied 12 years before sampling) decreased needle B concentrations in the four youngest needle age classes from 6–10 mg kg^-1 to 5 mg kg^-1. In boron fertilized plots the corresponding concentrations were 23–35 mg kg^-1 in control plots and 21–29 mg kg^-1 in limed plots. Both liming and B fertilizer decreased the Mn concentrations of needles. In the humus layer, total B concentration was increased by both lime and B fertilizer, and Ca and Mg concentrations and pH were still considerably higher in the limed plots than controls. Liming decreased the organic matter concentration in humus layer, whilst B fertilizer increased it.The results about B uptake were confirmed in a pot experiment, in which additionally the roles of increased soil pH and increased soil Ca concentration were separated by means of comparing the effects of CaCO₃ and CaSO₄. Two-year-old bare-rooted Norway spruce seedlings were grown in mor humus during the extension growth of the new shoot. The two doses of lime increased the pH of soil from 4.1 to 5.6 to 6.1, and correspondingly decreased the B concentrations in new needles from 22 to 12 to 9 mg kg^-1. However, CaSO₄ did not affect the pH of the soil or needle B concentrations. Hence the liming effect on boron availability in these soils appeared to be caused by the increased pH rather than increased calcium concentration.     

Fine root dynamics in a loblolly pine forest are influenced by free-air-CO2-enrichment: a six-year-minirhizotron study

Efforts to characterize carbon (C) cycling among atmosphere, forest canopy, and soil C pools are hindered by poorly quantified fine root dynamics. We characterized the influence of free‐air‐CO₂‐enrichment (ambient +200 ppm) on fine roots for a period of 6 years (Autumn 1998 through Autumn 2004) in an 18‐year‐old loblolly pine (Pinus taeda) plantation near Durham, NC, USA using minirhizotrons. Root production and mortality were synchronous processes that peaked most years during spring and early summer. Seasonality of fine root production and mortality was not influenced by atmospheric CO₂ availability. Averaged over all 6 years of the study, CO₂ enrichment increased average fine root standing crop (+23%), annual root length production (+25%), and annual root length mortality (+36%). Larger increase in mortality compared with production with CO₂ enrichment is explained by shorter average fine root lifespans in elevated plots (500 days) compared with controls (574 days). The effects of CO₂‐enrichment on fine root proliferation tended to shift from shallow (0–15 cm) to deeper soil depths (15–30) with increasing duration of the study. Diameters of fine roots were initially increased by CO₂‐enrichment but this effect diminished over time. Averaged over 6 years, annual fine root NPP was estimated to be 163 g dw m^?2 yr^?1 in CO₂‐enriched plots and 130 g dw m^?2 yr^?1 in control plots (P= 0.13) corresponding to an average annual additional input of fine root biomass to soil of 33 g m^?2 yr^?1 in CO₂‐enriched plots. A lack of consistent CO₂× year effects suggest that the positive effects of CO₂ enrichment on fine root growth persisted 6 years following minirhizotron tube installation (8 years following initiation of the CO₂ fumigation). Although CO₂‐enrichment contributed to extra flow of C into soil in this experiment, the magnitude of the effect was small suggesting only modest potential for fine root processes to directly contribute to soil C storage in south‐eastern pine forests.     

Does liquid fertilization affect fine root dynamics and lifespan of mycorrhizal short roots?

We studied effects of nitrogen, other nutrients and water (liquid fertilization; LF) on fine root dynamics (production, mortality) and life span of mycorrhizal short roots in a Norway spruce stand, using minirhizotrons. Data were collected and analyzed during a two-year period at depths of 0–20 cm, 21–40 cm and 41–85 cm, six years after the start of treatment. Relative to control (C), root production was lower in LF plots at depth 0–20 cm. Root production increased significantly at depth 41–85 cm. Fine root mortality in LF plots was higher at all depths. Life span of mycorrhizal short roots in LF plots was significantly lower than C plots and at the end of the study no mycorrhizal short roots were alive. It is suggested that the water and nitrogen input lower longevity of mycorrhizal short roots and promote fine root production at deeper soil layers.     

Effects of ammonium sulphate application on the rhizosphere,fine-root and needle chemistry in aPicea abies (L.) Karst. stand

Rhizosphere, fine-root and needle chemistry were investigated in a 28 year old Norway spruce stand in SW Sweden. The uptake and allocation pattern of plant nutrients and aluminium in control plots (C) and plots repeatedly treated with ammonium sulphate (NS) were compared. Treatments started in 1988. Current year needles, one-year-old needles and cylindrical core samples of the LFH-layer and the mineral soil layers were sampled in 1988, 1989 and 1990. Compared to the control plots, pH decreased significantly in the rhizosphere soil in the NS plots in 1989 and 1990 while the SO₄-S concentration increased significantly. Aluminium concentration in the rhizosphere soil was generally higher in the NS plots in all soil layers, except at 0–10 cm depths, both in 1989 and 1990. Calcium, Mg and K concentrations also increased after treatment with ammonium sulphate. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment significantly reduced Mg concentrations in fine roots in all layers in 1990. A similar trend was found in the needles. Ca concentrations in fine roots were significantly lower in the NS plots in the LFH layer in 1990 and the same pattern was found in the current needles. The N and S concentrations of both fine roots and needles were significantly higher in the NS plots. It was suggested that NS treatment resulted in displacement of Mg, Ca and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

Soil warming and CO2 enrichment induce biomass shifts in alpine tree line vegetation

Responses of alpine tree line ecosystems to increasing atmospheric CO₂ concentrations and global warming are poorly understood. We used an experiment at the Swiss tree line to investigate changes in vegetation biomass after 9 years of free air CO₂ enrichment (+200 ppm; 2001–2009) and 6 years of soil warming (+4 °C; 2007–2012). The study contained two key tree line species, Larix decidua and Pinus uncinata, both approximately 40 years old, growing in heath vegetation dominated by dwarf shrubs. In 2012, we harvested and measured biomass of all trees (including root systems), above‐ground understorey vegetation and fine roots. Overall, soil warming had clearer effects on plant biomass than CO₂ enrichment, and there were no interactive effects between treatments. Total plant biomass increased in warmed plots containing Pinus but not in those with Larix. This response was driven by changes in tree mass (+50%), which contributed an average of 84% (5.7 kg m^?2) of total plant mass. Pinus coarse root mass was especially enhanced by warming (+100%), yielding an increased root mass fraction. Elevated CO₂ led to an increased relative growth rate of Larix stem basal area but no change in the final biomass of either tree species. Total understorey above‐ground mass was not altered by soil warming or elevated CO₂. However, Vaccinium myrtillus mass increased with both treatments, graminoid mass declined with warming, and forb and nonvascular plant (moss and lichen) mass decreased with both treatments. Fine roots showed a substantial reduction under soil warming (?40% for all roots <2 mm in diameter at 0–20 cm soil depth) but no change with CO₂ enrichment. Our findings suggest that enhanced overall productivity and shifts in biomass allocation will occur at the tree line, particularly with global warming. However, individual species and functional groups will respond differently to these environmental changes, with consequences for ecosystem structure and functioning.     

Nitrogen transformations in limed and nitrogen fertilized soil in Norway spruce stands

Nitrogen transformations in the soil, and the resulting changes in carbon and nitrogen compounds in soil percolate water, were studied in two stands of Norway spruce (Picea abies L.). Over the last 30 years the stands were repeatedly limed (total 6000 kg ha^–1), fertilized with nitrogen (total about 900 kg ha^–1), or both treatments together. Both aerobic incubations of soil samples in the laboratory, and intact soil core incubations in the field showed that in control plots ammonification widely predominated over nitrification. In both experiments nitrogen addition increased the formation of mineral-N. In one experiment separate lime and nitrogen treatments increased nitrification, in the other, only lime and nitrogen addition together had this effect. In one experiment immobilization of nitrogen to soil microbial biomass was lower in soil only treated with nitrogen. Soil percolate water was collected by means of lysimeters placed under the humus layer and 10 cm below in the mineral soil. Total N, NH₄-N and NO₃-N were measured, and dissolved organic nitrogen was fractioned according to molecular weight. NO₃-N concentrations in percolate water, collected under the humus layer, were higher in plots treated with N-fertilizer, especially when lime was also added. The treatments had no effect on the N concentrations in mineral soil. A considerable proportion of nitrogen was leached in organic form.     

Patterns of restoration of soil physciochemical properties and microbial biomass in different landslide sites in the sal forest ecosystem of Nepal Himalaya

The pattern of natural restoration in soil components and processes was documented in five landslide-damaged (1–58-year-old) sites in the moist tropical sal (Shorea robusta) forest ecosystem of Nepal Himalaya. Comparisons were made with an undisturbed forest site in the same region. Concentrations of soil organic C, total N, total P and extractable nutrients (Ca, Mg and K) increased with the age of sites. The 58-year-old site showed concentrations of soil organic C, total N and total P that were 75–89% of concentrations in the undisturbed sal forest. The soil microbial biomass, the active fraction of soil organic matter, showed similar seasonal variations at all sites. The amount of mean microbial biomass (expressed as C, N and P contents) increased 4–5 times at the 58-year-old site relative to the 1-year-old site, and the bulk increase occurred within the initial 15 year. The increase in the C/N ratio of soil microbial biomass with age (9.4–11.6 years) reflected change in its composition. Although the net N-mineralization rate increased consistently until 58 years of age, the proportion of nitrification rate relative to ammonification rate distinctly decreased beyond 40 years. On the other hand, the soil available-N (both NO₃⁻ and NH₄⁺) concentrations increased from 1 to 40 year and then declined; with age the proportion of NH₄⁺ increased, however. Rates of restoration in soil properties were faster in the early successional stages (1–15 year) than late stages. Among different soil properties the restoration of soil microbial biomass (C and N) was faster than soil organic C and total N. Best fit power function models showed that the estimated times for the 58-year-old site to reach the level of the undisturbed, mature sal forest would be about 30–35 year for microbial biomass (C and N) and about 100–150 year for organic C and total N. Higher accumulation of soil microbial biomass and high N-mineralization rate at late successional stages indicated the re-establishment of enriched soil and restitution of nutrient cycling during the course of ecosystem restoration.     

Needle chemistry in young Norway spruce stands after application of crushed wood ash

Nutrient concentrations in current and 1-year-old needles were analyzed annually for 5 years after application of hardened wood ash in 1–4-year-old Norway spruce (Picea abies (L.) Karst.) stands within a range of climate and fertility gradients. At each site, 3000 kg ha^–1 hardened wood ash of two types, Nymölla and Perstorp, was applied in a randomized block design. Wood ash Nymölla contained 12 kg ha^–1 P, 30 kg ha^–1 K, 891 kg ha^–1 Ca, 72 kg ha^–1 Mg and wood ash Perstorp contained 12 kg ha^–1 P, 60 kg ha^–1 K, 486 kg ha^–1 Ca, and 60 kg ha^–1 Mg. The ash was intended to compensate for nutrients removed at the preceding harvest when logging residues were collected and removed from the site (whole-tree harvesting). The climate gradient included four climate zones throughout Sweden and each of these included a fertility gradient of three sites classified according to their ground vegetation type. There were no effects on nutrient concentrations in the needles 1 year after the application of wood ash. Five years after ash application, the concentrations of P, K and Ca in current and 1-year-old needles were higher than in the control plots. The results were consistent over all stands, irrespective of climate zone and fertility status. P and K concentrations were higher in spruce needles from plots treated with Perstorp wood ash, whereas Ca concentrations were higher in those of Nymölla treated plots. Analyses across all study sites revealed a treatment effect in terms of increased ratios of P:N, K:N and Ca:N in 1-year-old needles. The ratio P:N tended to increase with time in the Perstorp wood ash treatment compared with the control. The needle concentrations of Mg and S were not affected by the ash applications. The increase in needle nutrient concentrations after application of hardened wood ash suggests that wood ash recycling could be used in order to replace nutrients removed at whole-tree harvesting.     

Fine root production and nutrient content in wet and moist arctic tundras as influenced by chronic fertilization

We used ingrowth cores to estimate fine root production in organic soils of wet sedge and moist tundra ecosystems near Toolik Lake on Alaska's North Slope. Root-free soil cores contained in nylon mesh tubes (5 cm diameter, 20–30 cm long) were placed in control and chronically fertilized (N plus P) plots in mid-August 1994 and were retrieved 1 year later. Estimated fine root production in control plots was 75 g m^–2 year^–1 in wet sedge and 56 g m^–2 year^–1 in moist tussock tundra. Fine root production in fertilized plots was 85 g m^–2 year^–1 in wet sedge and 67 g m^–2 year^–1 in moist tussock tundra. Although our estimates of fine root production were higher on fertilized than control plots, differences were not statistically significant within either tundra type. Comparisons between our estimates of fine root production and other estimates of aboveground (plus rhizome) production on the same (wet sedge tundra) or similar (moist tussock tundra) plots suggest that fine root production was about one-third of total net primary production (NPP) under non-fertilized conditions and about one-fifth of total NPP under chronic fertilization. Fine root N and P concentrations increased with fertilization in both tundra types, but P concentrations increased more than N concentrations in wet sedge tundra, whereas relative increases in N and P concentrations in moist tundra roots were similar. These data are consistent with other studies suggesting that NPP in wet sedge tundra is often P limited and that co-limitation by N and P is more important in moist tussock tundra.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.     

Tree growth and soil acidification in response to 30 years of experimental nitrogen loading on boreal forest

Relations among nitrogen load, soil acidification and forest growth have been evaluated based on short‐term (<15 years) experiments, or on surveys across gradients of N deposition that may also include variations in edaphic conditions and other pollutants, which confound the interpretation of effects of N per se. We report effects on trees and soils in a uniquely long‐term (30 years) experiment with annual N loading on an un‐polluted boreal forest. Ammonium nitrate was added to replicated (N=3) 0.09 ha plots at two doses, N1 and N2, 34 and 68 kg N ha^?1 yr^?1, respectively. A third treatment, N3, 108 kg N ha^?1 yr^?1, was terminated after 20 years, allowing assessment of recovery during 10 years. Tree growth initially responded positively to all N treatments, but the longer term response was highly rate dependent with no gain in N3, a gain of 50 m³ ha^?1 stemwood in N2 and a gain of 100 m³ ha^?1 stemwood in excess of the control (N0) in N1. High N treatments caused losses of up to 70% of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺) in the mineral soil, along with decreases in pH and increases in exchangeable Al³⁺. In contrast, the organic mor‐layer (forest floor) in the N‐treated plots had similar amounts per hectare of exchangeable base cations as in the N0 treatment. Magnesium was even higher in the mor of N‐treated plots, providing evidence of up‐lift by the trees from the mineral soil. Tree growth did not correlate with the soil Ca/Al ratio (a suggested predictor of effects of soil acidity on tree growth). A boron deficiency occurred on N‐treated plots, but was corrected at an early stage. Extractable NH₄⁺ and NO₃^?were high in mor and mineral soils of on‐going N treatments, while NH₄+ was elevated in the mor only in N3 plots. Ten years after termination of N addition in the N3 treatment, the pH had increased significantly in the mineral soil; there were also tendencies of higher soil base status and concentrations of base cations in the foliage. Our data suggest the recovery of soil chemical properties, notably pH, may be quicker after removal of the N‐load than predicted. Our long‐term experiment demonstrated the fundamental importance of the rate of N application relative to the total amount of N applied, in particular with regard to tree growth and C sequestration. Hence, experiments adding high doses of N over short periods do not mimic the long‐term effects of N deposition at lower rates.     

Role of plant parasitic nematodes and abiotic soil factors in growth heterogeneity of sugarcane on a sandy soil in South Africa

A feature of many sugarcane fields is the patchy growth. This is often thought to be due to physical or chemical differences in the soil. In this paper we investigate the causes of growth heterogeneity of sugarcane on a sandy soil in KwaZulu-Natal, South Africa. To identify the factors that were associated with the good and poor areas, soil texture, pH, organic matter content and a number of soil chemical elements and nematode community data were subjected to principal component analysis (PCA). The numbers of each of the nematode species (Meloidogyne sp., Pratylenchus zeae, Helicotylenchus dihystera, Xiphinema elongatum and Paratrichodorus sp.) were first converted to relative proportions of ectoparasites and endoparasites. The data were collected from the 2nd ratoon crop of a nematicide trial, where half of the plots had been treated with aldicarb in the preceding two crops. Yields of control plots varied from 34.7 to 126.8 t cane ha^–1 and from 85.4 to 138.7 for the treated plots. The yield data were centred and normalised separately for the treated and control plots and the values projected on the trial map to study spatial distribution. Plots with above-average yields, whether treated or untreated, occurred in the lower part of the trial site. The PCA factorial values were also projected onto the map of the trial. According to the first factor of the analysis of the abiotic soil characteristics in the 0–20 cm surface layer, the trial site could be divided into two areas, one on the left and one on the right. PCA of the soil data from the 0–20 and 20–40 cm layers showed that there was little or no difference between the two that might explain the two growth areas. However, analysis of the nematode community distinguished two main areas that largely corresponded to the distribution of the plots of low and high yielding cane. Correlation analysis confirmed the relationship between nematodes and yield. H. dihystera was positively correlated with yield of cane whereas the reverse was true for the Meloidogyne species.     

Tree species selection and soil tillage in alley cropping systems with Phaseolus vulgaris L. in a humid premontane climate: biomass production,nutrient cycling and crop responses

The development of alley cropping systems is based on the assumption that leguminous trees planted in hedgerows influence the yield of associated crops favourably by means of the additional nutrient pool applied to the soil through tree prunings. An on-station field study (split-plot design in a randomised block design) was conducted on an Eutric Cambisol under humid premontane climate conditions in Costa Rica in order to evaluate the ability of Erythrina poeppigiana, Calliandra calothyrsus and Gliricidia sepium to increase bean (Phaseolus vulgaris) yields compared to sole cropping. Soil tillage was applied as a sub-treatment in order to evaluate if soil preparation would additionally alter soil fertility and bean yield. After seven years with pruning twice per year, the size of both the total N and P pool in the pruned tree material was about three times higher for Erythrina prunings than for Calliandra and Gliricidia prunings. Two and five weeks after mulch application 50–150% higher inorganic N pools were measured in the soil from Erythrina plots, the bean shoot biomass at harvest was increased by 65–100% and the bean yield was 15–50% higher than in plots with beans alone. Hence, of the three tree species, Erythrina was the best choice for alley cropping systems in the pedoclimatic environment studied. Soil tillage reduced bean yield, soil organic matter, total soil N content and soil microbial biomass N in the top soil and is not recommended for similar soils in humid premontane climates.     

The influence of drainage and soil phosphorus on the vegetation of Douala-Edea Forest Reserve,Cameroun

All living trees (30 cm gbh) were enumerated in 104 80×80 m plots arranged along four transects in the Douala-Edea Forest Reserve Cameroun, a system of low-lying ancient coastal sand dunes interspersed by numerous streams and swamps. The extent of permanent and seasonal swamps was recorded for each plot. Two hundred thirty taxa were recognized of which 63% were identified to species. Mean tree density was 376 ha^–1, basal area 31.0 m² ha^–1 and number of species per plot 39. The Olacaceae were the most abundant family in terms of basal area, but the Euphorbiaceae the most frequently represented. The most abundant species wasCoula edulis (Olacaceae). Twenty-two plots had most of their area permanently or seasonally swamped. Percentage sand, silt and clay ranged between 32–100, 0–64, 0–21% respectively. The ranges for other variables recorded were: pH (2.7–5.4), organic carbon (1.5–12.4%), available phosphorus (7–90 ppm) and potassium (28–188 ppm), and nitrogen (ammonium 4–40 ppm, nitrate 1–12 ppm).Classification of the plots on the basis of six soil variables provided three large distinct groups: swamp plots and non-swamp plots, the latter divided into plots of low and high available soil phosphorus. Swamp plots were distinguished by high abundances ofProtomegabaria stapfiana andLibrevillea klainei, though correspondence ordination of plots in these groups showedP. stapfiana associated with more clayey soils andLibrevillea klainei (andGluema ivorensis) on the very sandy soils. Direct gradient analysis highlighted several species associated with these lower phosphorus soils. Available soil phosphorus is not as low at Douala-Edea as in parts of Korup, and the association of these Douala-Edea soils with the Caesalpinioideae is correspondingly weaker.Nomenclature follows Aubréville (1963–1983).The field work was supported by grant numbers RR00167-14, RR00167-15 and RR0167-16 from the National Institutes of Health for the operation of the Wisconsin Regional Primate Research Center, and N.A.T.O. Scientific Affairs grant number 1748 (to PGW and JSG). It was greatly facilitated by the skill and dedication of Ferdinand Namata. R. M. Polhill and D. W. Thomas assisted considerably in the identification of plant species. Sue Gartlan collected and collated the meteorological data, besides other field support. In the field phase J.S.G., P.G.W. and D.B.McK. were researches attached to the National Office of Scientific and Technical Research (ONAREST), Yaoundé. We thank M. D. Swaine for comments on earlier drafts, R. Letouzey for checking species nomenclature, the Computer Unit at Stirling University for facilities, M. Burnett for the typing at Stirling, and the Department of Soil Science, University of Wisconsin, for undertaking the soil chemical analyses.Reprint requests to D.McC.N. at Stirling.Publication No. 23-025 of the Wisconsin Regional Primate Center.     

Destructive and non-destructive measurements of residual crop residue and phosphorus effects on growth and composition of herbaceous fallow species in the Sahel

Little is known about the residual effects of crop residue (CR) and phosphorus (P) application on the fallow vegetation following repeated cultivation of pearl millet [Pennisetum glaucum (L.) R. Br.] in the Sahel. The objective of this study, therefore, was (i) to measure residual effects of CR, mulched at annual rates of 0, 500, 1000 and 2000 kg CR ha^–1, broadcast P at 0 and 13 kg P ha^–1 and P placement at 0, 1, 3, 5 and 7 kg P ha^–1 on the herbaceous dry matter (HDM) 2 years after the end of the experiment and (ii) to test a remote sensing method for the quantitative estimation of HDM. Compared with unmulched plots, a doubling of HDM was measured in plots that had received at least 500 kg CR ha^–1. Previous broadcast P application led to HDM increases of 14% compared with unfertilised control plots, whereas no residual effects of P placement were detected. Crop residue and P treatments caused significant shifts in flora composition. Digital analysis of colour photographs taken of the fallow vegetation and the bare soil revealed that the number of normalised green band pixels averaged per plot was highly correlated with HDM (r = 0.86) and that red band pixels were related to differences in soil surface crusting. Given the traditional use of fallow vegetation as fodder, the results strongly suggest that for the integrated farming systems of the West African Sahel, residual effects of soil amendments on the fallow vegetation should be included in any comprehensive analysis of treatment effects on the agro-pastoral system.