Effects of frequent fires and grazing on stable nitrogen isotope ratios of vegetation in northern Australia

The ratios of stable nitrogen isotopes expressed as δ¹⁵N values can indicate the openness of nitrogen cycles in ecosystems. Southwards through the Northern Territory, values of foliar δ¹⁵N in savanna trees increase as mean annual rainfall decreases from approximately 1800 mm to approximately 750 mm, with foliar δ¹⁵N thereafter decreasing toward arid central Australia. Recent literature argues that this pattern is caused by higher grazing intensity in semi‐arid savannas, but counter views have attributed the pattern more directly to variations in aridity. In this paper, grazed and ungrazed sites in a semi‐arid savanna are compared, and it is shown that grazing has a relatively small effect on the positive foliar δ¹⁵N values of grasses, but no effect on δ¹⁵N values of trees. This gives little support to the argument that variations in grazing pressure at the scale of hundreds of kilometres could result in detectable differences in the foliar δ¹⁵N values of trees. I then compare the semi‐arid savannas with mesic savannas, where fires are frequent, and with mesic rainforests, which are rarely burnt. Greater foliar δ¹⁵N values in rainforest and fire‐excluded mesic savannas than in frequently burnt savannas suggests that fire regimes affect foliar δ¹⁵N. The previously observed pattern in δ¹⁵N values along the rainfall gradient in the Northern Territory is consistent with trends in fire frequency and possible direct effects of fire, but further work is required to determine the relative impacts of aridity and fire. Within a particular rainfall regime, foliar δ¹⁵N values may indicate historical fire frequencies.     

Natural 15N abundance of vegetation and soil in the Kapalga savanna,Australia

Abstract The natural abundance of the stable isotope ¹⁵N was measured in different vegetation components and in the soil of a northern Australian savanna. Most of the vegetation was found to be ¹⁵N-depleted compared to atmospheric N₂. Herbaceous legumes, perennial grasses, tree legumes, non-legume trees and annual grasses exhibited mean δ¹⁵N of ? 1.7, ? 0.8, ? 0.7, 0.0 and + 0.3‰, respectively. These results are in good agreement with previous studies. Legumes exhibit slightly negative values, indicating that they are likely to be nitrogen-fixing plants. Non-legume plants have a δ¹⁵N close to zero, which could equally result from non-symbiotic fixation, soil organic matter mineralization, or fresh root litter mineralization. In contrast, soil organic matter was ¹⁵N-enriched. Values of δ¹⁵N increased with depth and were + 2.5, + 5.2 and +6.1‰ in the 0–10, 10–20 and 20–40cm layers, respectively. Soil organic matter δ¹⁵N shows a typical profile of mature soils.     

Spatial and temporal soil moisture resource partitioning by trees and grasses in a temperate savanna, Arizona, USA

Stable isotope analysis was used to determine sources of water used by coexisting trees and grasses in a temperate savanna dominated by Quercus emoryi Torr. We predicted that (1) tree seedlings and bunchgrasses utilize shallow sources of soil water, (2) mature savanna trees use deeper sources of water, and (3) trees switch from shallow to deep water sources within 1 year of germination. We found that Q. emoryi trees, saplings, and seedlings (about 2 months, 1 year, and 2 years old), and the dominant bunchgrass [Trachypogon montufari (H.B.K.) Nees.] utilized seasonally available moisture from different depths within the soil profile depending on size/age relationships. Sapling and mature Q. emoryi acquired water from >50 cm deep, 2-month-old seedlings utilized water from <15 cm, and 1- and 2-year-old seedlings and grasses used water from between 20 cm and 35 cm. This suggests that very young seedlings are decoupled from grasses in this system, which may facilitate germination and early establishment of Q. emoryi within extant stands of native grasses. The potential for subsequent interaction between Q. emoryi and native grasses was evidenced by similar patterns of soil water use by 1- and 2-year-old seedlings and grasses. Q. emoryi seedlings did not switch from shallow to deep sources of soil water within 2 years of germination: water use by these seedlings apparently becomes independent of water use by grasses after 2 years of age. Finally, older trees (saplings, mature trees) use water from deeper soil layers than grasses, which may facilitate the stable coexistence of mature trees and grasses. Potential shifts in the seasonality of precipitation may alter interactions between woody plants and grasses within temperate savannas characterized by bimodal precipitation regimes: reductions in summer precipitation or soil moisture may be particularly detrimental to warm-season grasses and seedlings of Q. emoryi. Received: 21 November 1996 / Accepted: 2 May 1997     

Multiyear fate of a 15N tracer in a mixed deciduous forest: retention,redistribution, and differences by mycorrhizal association

The impact of atmospheric nitrogen deposition on forest ecosystems depends in large part on its fate. Past tracer studies show that litter and soils dominate the short‐term fate of added ¹⁵N, yet few have examined its longer term dynamics or differences among forest types. This study examined the fate of a ¹⁵N‐ tracer over 5–6 years in a mixed deciduous stand that was evenly composed of trees with ectomycorrhizal and arbuscular mycorrhizal associations. The tracer was expected to slowly mineralize from its main initial fate in litter and surface soil, with some ¹⁵N moving to trees, some to deeper soil, and some net losses. Recovery of added ¹⁵N in trees and litterfall totaled 11.3% both 1 and 5–6 years after the tracer addition, as ¹⁵N redistributed from fine and especially coarse roots into cumulative litterfall and small accumulations in woody tissues. Estimates of potential carbon sequestration from tree ¹⁵N recovery amounted to 12–14 kg C per kg of N deposition. Tree ¹⁵N acquisition occurred within the first year after the tracer addition, with no subsequent additional net transfer of ¹⁵N from detrital to plant pools. In both years, ectomycorrhizal trees gained 50% more of the tracer than did trees with arbuscular mycorrhizae. Much of the ¹⁵N recovered in wood occurred in tree rings formed prior to the ¹⁵N addition, demonstrating the mobility of N in wood. Tracer recovery rapidly decreased over time in surface litter material and accumulated in both shallow and deep soil, perhaps through mixing by earthworms. Overall, results showed redistribution of tracer ¹⁵N through trees and surface soils without any losses, as whole‐ecosystem recovery remained constant between 1 and 5–6 years at 70% of the ¹⁵N addition. These results demonstrate the persistent ecosystem retention of N deposition even as it redistributes, without additional plant uptake over this timescale.     

Increases of Soil C, N, and P Pools Along an Acacia Tree Density Gradient and Their Effects on Trees and Grasses

Nitrogen (N) fixing trees including many species of Acacia are an important though variable component of savanna ecosystems. It is known that these trees enrich the soil with carbon (C) and N, but their effect on the combined C:N:P stoichiometry in soil is less well understood. Theory suggests that they might reduce available phosphorus (P), creating a shift from more N-limited conditions in grass-dominated to more P-limited conditions in tree-dominated sites, which in turn could feed back negatively on the trees’ capacity to fix N. We studied the effects of Acacia zanzibarica tree density upon soil and foliar N:P stoichiometry, and the N₂-fixation rates of trees and leguminous herbs in a humid Tanzanian savanna. Foliar N:P ratios and N₂-fixation rates of trees remained constant across the density gradient, whereas soil C, N and organic P pools increased. In contrast, the N:P ratio of grasses increased and N₂-fixation rates of leguminous herbs decreased with increasing tree density, indicating a shift towards more P-limited conditions for the understory vegetation. These contrasting responses suggest that trees and grasses have access to different sources of N and P, with trees being able to access P from deeper soil layers and perhaps also utilizing organic forms more efficiently.     

Sources of increased N uptake in forest trees growing under elevated CO2: results of a large‐scale 15N study

Nitrogen availability in terrestrial ecosystems strongly influences plant productivity and nutrient cycling in response to increasing atmospheric carbon dioxide (CO₂). Elevated CO₂ has consistently stimulated forest productivity at the Duke Forest free‐air CO₂ enrichment experiment throughout the decade‐long experiment. It remains unclear how the N cycle has changed with elevated CO₂ to support this increased productivity. Using natural‐abundance measures of N isotopes together with an ecosystem‐scale ¹⁵N tracer experiment, we quantified the cycling of ¹⁵N in plant and soil pools under ambient and elevated CO₂ over three growing seasons to determine how elevated CO₂ changed N cycling between plants, soil, and microorganisms. After measuring natural‐abundance ¹⁵N differences in ambient and CO₂‐fumigated plots, we applied inorganic ¹⁵N tracers and quantified the redistribution of ¹⁵N for three subsequent growing seasons. The natural abundance of leaf litter was enriched under elevated compared to ambient CO₂, consistent with deeper rooting and enhanced N mineralization. After tracer application, ¹⁵N was initially retained in the organic and mineral soil horizons. Recovery of ¹⁵N in plant biomass was 3.5 ± 0.5% in the canopy, 1.7 ± 0.2% in roots and 1.7 ± 0.2% in branches. After two growing seasons, ¹⁵N recoveries in biomass and soil pools were not significantly different between CO₂ treatments, despite greater total N uptake under elevated CO₂. After the third growing season, ¹⁵N recovery in trees was significantly higher in elevated compared to ambient CO₂. Natural‐abundance ¹⁵N and tracer results, taken together, suggest that trees growing under elevated CO₂ acquired additional soil N resources to support increased plant growth. Our study provides an integrated understanding of elevated CO₂ effects on N cycling in the Duke Forest and provides a basis for inferring how C and N cycling in this forest may respond to elevated CO₂ beyond the decadal time scale.     

Can stable carbon isotopes (δ13C) in soil carbon be used to describe the dynamics of Eucalyptus savanna–rainforest boundaries in the Australian monsoon tropics?

The history of isolated patches of monsoon rainforest within large tracts of Eucalyptus savanna is poorly understood because of the scarcity of reliable palaeoecological records in the Australian monsoon tropics. Elsewhere in the world, the ratio of the stable isotopes ¹³C to ¹²C (δ¹³C) in soil organic matter has shed light on the dynamics of rainforest–savanna boundaries because tropical grasses with the C4 photosynthetic pathway have a distinct δ¹³C signature (–17 to –9‰) compared with that of woody plants with the C3 photosynthetic pathway (–32 to –22‰). In order to determine the magnitude of the variation in δ¹³C, unreplicated soil profiles were sampled beneath different vegetation types on three boundaries between Eucalyptus savanna and rainforest that were both growing on Tertiary age laterite parent material. Replicated (n = 3) soil profiles, which were also derived from Tertiary age laterite, were sampled from beneath: (i) dense stands of African grasses within a frequently burnt Eucalyptus savanna; and within the same long unburnt Eucalyptus savanna, (ii) patches of African and natives grasses and (iii) clumps of Acacia trees. The strongly negative δ¹³C values of soil organic matter derived from the frequently burnt and long unburnt grassy understoreys in the Eucalyptus savannas showed that a considerable amount of the soil carbon was derived from C3 (woody) species despite the presence of a ground layer dominated by C4 grasses. However, a feature of these data was the considerable variability among the three ‘replicate’ profiles. The surface soil samples from beneath three clumps of Acacia trees in the unburnt Eucalyptus savanna had much less variable δ¹³C values and were similar to two of the three monsoon rainforests sampled. The pattern of δ¹³C values from unreplicated soil profiles from different vegetation types across three rainforest boundaries was also very variable and not always obviously related the known disturbance history of the extant vegetation. Given the considerable variability within and between vegetation types with contrasting disturbance histories, it is concluded that the use of carbon stable isotopes to advance understanding of the dynamics of rainforest and Eucalyptus savanna boundaries will require further development, such as determination of the ¹⁴C age and δ¹³C values of different soil carbon fractions.     

Stable isotopes analysis to assess the trophic role of ants in a Mediterranean agroecosystem

1 Stable isotopes signatures (δ¹³C and δ¹⁵N) of the most important tree‐dwelling ants in an olive orchard were examined, together with the signatures of the most common herbivores, predators and sap‐sucking insects. The olive orchard consists of separate subunits (trees) surrounded by a matrix of grasses or bare ground, and the role of ants in such a system is not fully understood. 2 None of the selected ant species was exclusive to the olive trees because they were also observed foraging on vegetation (mainly thistle) under the tree crowns. Hence, the relative contributions of these two sources of energy (olive trees versus herbs/grasses) were assessed by comparing the δ¹³C of ants with the signatures of plants and those of other arthropods collected on the trees and on nearby thistles. Next, the trophic level occupied by the ants and their ecological role within the olive food web were determined by examining the δ¹⁵N values and their relationship with indices of ecological performance measuring the potential pressure exerted by each species on the ecosystem. 3 The analysis of ¹³C signatures revealed a different contribution of the two energy sources, olive trees versus herbs and grasses, with the former being more important for ants. The analysis of ¹⁵N signatures suggested separate roles for different ant species: some (Crematogaster scutellaris, Lasius lasioides) occupied a higher trophic level, mostly involved in predation, whereas others (Camponotus piceus, Camponotus lateralis) occupied a lower level, probably involved more in homopteran tending. A fifth species (Camponotus aethiops) was in an intermediate position. Finally, the δ¹⁵N levels of the species were significantly correlated with indices of ecological performance.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

The distribution of tree and grass roots in savannas in relation to soil nitrogen and water

Here we describe the fine root distribution of trees and grasses relative to soil nitrogen and water profiles. The primary objective is to improve our understanding of edaphic processes influencing the relative abundance of trees and grasses in savanna systems. We do this at both a mesic (737 mm MAP) site on sandy-loam soils and at an arid (547 mm MAP) site on clay rich soils in the Kruger National Park in South Africa. The proportion of tree and grass fine roots at each soil depth were estimated using the δ¹³C values of fine roots and the δ¹³C end members of the fine roots of the dominant trees and grasses at our study sites. Changes in soil nitrogen concentrations with depth were indexed using total soil nitrogen concentrations and soil δ¹⁵N values. Soil water content was measured at different depths using capacitance probes. We show that most tree and grass roots are located in the upper layers of the soil and that both tree and grass roots are present at the bottom of the profile. We demonstrate that root density is positively related to the distribution of soil nitrogen and negatively related to soil moisture. We attribute the negative correlation with soil moisture to evaporation from the soil surface and uptake by roots. Our data is a snapshot of a dynamic process, here the picture it provides is potentially misleading. To understand whether roots in this system are primarily foraging for water or for nitrogen future studies need to include a dynamic component.     

Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree–grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C₄ grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C₄ grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C₄ grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.     

Resilience and Thresholds in Savannas: Nitrogen and Fire as Drivers and Responders of Vegetation Transition

Resilience theory suggests that ecosystems can persist for long periods, before changing rapidly to a new vegetation phase. Transition between phases occurs when ecological thresholds have been crossed, and is followed by a reorganization of biotic and environmental interactions, leading to the emergence of a new vegetation phase or quasi-stable state. Savannas are dynamic, complex systems in which fire, herbivory, water and nutrient availability interact to determine tree abundance. Phase and transition has been observed in savannas, but the role of these different possible drivers is not always clear. In this study, our objectives were to identify phase and transition in the fossil pollen record, and then to explore the role of nitrogen and fire in these transitions using δ¹⁵N isotopes and charcoal abundance. We present palaeoenvironmental data from the Kruger National Park, South Africa, which show transition between grassland and savanna phases. Our results show transition at the end of the ninth century A.D. from a nutrient- and herbivore-limited grazing lawn, in which fire was absent and C₄ grasses were the dominant and competitively superior plant form, to a water-, fire- and herbivory-limited semi-arid savanna, in which C₄ grasses and C₃ trees and shrubs co-existed. The data accord with theoretical frameworks that predict that variability in ecosystems clusters in regions of higher probability space, interspersed by rapid transitions between these phases. The data are also consistent with the idea that phase transitions involve switching between different dominant driving processes or limiting factors.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

Topsoil translocation for Brazilian savanna restoration: propagation of herbs,shrubs, and trees

Topsoil translocation has been used for vegetation restoration throughout the world, but it has been poorly tested within savannas. This study describes Brazilian savanna (cerrado) regeneration for the first 3 years following topsoil translocation. The topsoil was stripped from 2.5 ha of savanna and spread on 1 ha of an abandoned laterite quarry in the Federal District, Brazil. We assessed vegetation structure and species composition in 18 circular plots (3.14/m²) after 5 and 15 months and in 30 circular plots after 37 months. In the last floristic survey, the coverage of herbs was estimated using the step‐point method. To verify the source of regeneration, a total of 181 shrubs and trees were excavated over the first 2 surveys. After 3 years, 24, 40, and 21 species of herbs, shrubs, and trees, respectively, had been recorded by the surveys. Of the 33 families found, Fabaceae, Poaceae, and Asteraceae were the most representative. At 5 and 15 months, 91 and 83% of the individuals (shrubs and trees combined) were derived from resprouting, respectively. Shrub and tree stem density reached 3.2/m² at 5 months, but declined to 0.5/m² at 37 months. By the final survey, native and exotic grasses completely covered the ground. Topsoil translocation was effective for the propagation of native herbs, shrubs, and trees, despite the need to control invasive grasses. The large number of shrub and tree resprouts from roots suggests that the bud bank is an important component of the topsoil for savanna restoration.     

Not all forests are expanding over central Brazilian savannas

Recently we reported on the expansion of riparian forests into savannas in central Brazil. To enlarge the scope of the earlier study we investigated whether upland deciduous and xeromorphic forests behaved similarly. We investigated past vegetation changes that occurred in forest/savanna transitions using carbon isotope ratios (δ¹³C) measured in the soil organic matter as a tracer. We analyzed the ¹⁴C activity where δ¹³C showed major shifts in vegetation. The role of soil chemical and physical attributes in defining vegetation distribution is discussed. Structural changes in vegetation were found to be associated with shifts in the isotope composition (δ¹³C) of soil organic matter. This was attributed to intrinsic differences in the biomass of trees and grasses and allowed for the determination of past shifts in vegetation by evaluating δ¹³C at different depths. The deciduous forest decreased in area approximately 980 years ago. Tree cover increased in the xeromorphic forest, but the border stayed stable through time. The deciduous forest and adjacent savanna have eutrophic soils while the xeromorphic forest and adjacent savanna have dystrophic soils. However, greater organic carbon, nitrogen and phosphorus concentrations are observed in the forests. We provide concrete evidence of deciduous forest retreat unlike the stability observed in the xeromorphic forest/savanna boundary. These results contrast with the expansion of riparian forests recently reported in the same region.     

Partitioning of 15N-labeled ammonium and nitrate among soil, litter, below- and above-ground biomass of trees and understory in a 15-year-old Picea abies plantation

The partitioning of nitrogen deposition among soil, litter, below- and above-ground biomass of trees and understory vegetation was investigated in a 15-year-old Picea abies (L.) Karst. plantation in the Fichtelgebirge, Germany, by labeling with 62 mg of¹⁵N tracer per square meter in March 1991. Ammonium and nitrate depositions were simulated on five plots each, by labeling with either¹⁵N-NH₄ ⁺ or¹⁵N-NO₃ ^–, and the¹⁵N pulse was followed during two successive growing seasons (1991 and 1992). Total recovery rates of the¹⁵N tracer in the entire stand ranged between 93 and 102% for both nitrogen forms in 1991, and 82% in June 1992. ⁵ N ratios increased rapidly in all compartments of the ecosystem. Roots and soils (to 65 cm depth) showed significant¹⁵N enrichments for both¹⁵N-treatments compared to reference plots. Newly grown spruce tissues were more enriched than older ones, but the most enriched ¹⁵N values were found in the understory vegetation. Although spruce trees were a much larger pool (1860 g biomass/m²) than understory vegetation (Vaccinium myrtillus 333 g/m², Calluna vulgaris 142 g/m², Deschampsia flexuosa 22 g/m²), the ericaceous shrubs and the perennial grass were a much greater sink for the¹⁵N label. Eight months after labeling, 9% of the ammonium and 15% of the nitrate label were found in the understory. P.abies retained only 3% of the¹⁵N-ammonium and 7% of the¹⁵N-nitrate. The main sink for both¹⁵N tracers was the soil, where 87% of the ammonium and 79% of the nitrate tracer were found. The organic soil horizon (5-0 cm depth) contained 63% of the¹⁵N-ammonium and 46% of the¹⁵N -nitrate suggesting strong immobilization by microorganisms of both N forms. Eight months after tracer application, about 16% of both¹⁵N-tracers was found below 25 cm soil depth. This 16% corresponds well to a 20% decrease in the recovery of both¹⁵N tracers after 15 months and indicates a total loss out of the ecosystem. Highly enriched ¹⁵N values were found in fruit bodies of fungi growing in reference lots (no¹⁵N addition), although soils did not show increased ¹⁵N ratios. No transfer of¹⁵N-tracer between fungi and spruce or understory vegetation was apparent yet.     

Termitaria vs. mistletoe: Effects on soil properties and plant structure in a semi-arid savanna

Termite mounds by creating patches of increased resource availability (e.g. water and nutrients) are a major source of spatial heterogeneity in savannas. Likewise, mistletoes via input of nutrient-rich litter alter nutrient and water availability increasing environmental heterogeneity in semi-arid savanna. Despite this recognition, the influence of termitaria and mistletoe on soil properties and plant community have not been investigated together. We established eight 100 m² plots each on termitaria, under mistletoe-infected trees and in the surrounding savanna and examined the soil properties and the structure of Securinega virosa (Euphorbiaceae) and Euclea divinorum (Ebenaceae) in semi-arid savanna, southwest Zimbabwe. Soil properties significantly differed among the sampling sites (p = 0.001) with soils of increasing clay, soil moisture, pH and phosphorus, calcium and ammonium concentrations occurring on termite mounds. Soils under mistletoe-infected trees were associated with silt, organic matter, sodium, potassium, magnesium and nitrate and the surrounding savanna was associated with soils of increasing sand content. Plant structure also differed significantly between sites with greater basal area of both S. virosa and E. divinorum on termitaria relative to mistletoe-infected trees and the surrounding savanna. However, the stem density of S. virosa was greater under mistletoe-infected trees than on termitaria and in the surrounding savanna. Plant structural variables of individuals of the same species were affected by different soil properties across treatments. The major patterns showed that plant structure was influenced positively by soil moisture and nitrate and negatively by phosphorus on termitaria; positively by clay, soil moisture and ammonium and negatively by potassium under mistletoe-infected trees; and by phosphorus and calcium in the surrounding savanna. These findings show that soil properties, plant structure and their relationships differ between termitaria, mistletoe-infected trees and surrounding savanna, and these differences are suggested to increase heterogeneity in soil resources availability and vegetation structure in semi-arid savanna.     

Wood ash treatment affects seasonal N fluctuations in needles of adult<Emphasis Type="Italic"> Picea abies</Emphasis> trees: a <Superscript>15</Superscript>N-tracer study

A ¹⁵N-tracer experiment was carried out in a stand of adult spruce trees [Picea abies (L.) Karst.] located on the Swiss Plateau in order to investigate the effects of wood ash treatment on seasonal nitrogen fluctuations in fine roots and needles. Treatments included irrigation (W), liquid fertilization (LF) and wood ash (A) application. ¹⁵N fluctuation in fine roots and current to 3-year-old needles was studied after one ¹⁵N pulse for 2 consecutive years (1999, 2000). ¹⁵N tracer was rapidly incorporated into the fine roots of adult trees, and ¹⁵N values reached similar levels in all treatments 2 months after the pulse. In the needles, the largest increase in ¹⁵N was observed in those of the current year. Following the initial peak during spring growth, ¹⁵N values in needles of control trees showed an oscillating pattern through the season. This oscillation is attributed to the increased use of internal N sources, as soon as the roots can no longer meet the increased N demand during the sprouting phase. However, W-, LF- and A-treated trees no longer showed the oscillation in ¹⁵N. Additional water (W and LF) as well as fertilizer (A and LF) may have induced shifts in the microbial flora, thus increasing the unlabelled N release from the soil. The strongest dampening was observed for the A treatment, indicating sufficient N availability from the soil, and making intensive use of the internal N sources unnecessary. Treatment with wood ash thus resulted in a similar fertilizer response to liquid fertilization.     

Contrasting impacts of grass species on nitrogen cycling in a grazed Sudanian savanna

We investigated the impact of perennial and annuals grass species on nitrogen cycling in a Sudanian savanna of Burkina Faso. We also analysed how the local context in terms of grazing and soil properties modifies these impacts. We selected four plots differing both by the intensity of grazing by cattle and soil depth, and used soil and grass biomass ¹⁵N as integrative indicators of N cycle. If perennials are able to foster a more efficient nitrogen cycling there should be lower ¹⁵N abundances in their biomass and soil. If soil depth and cattle pressure significantly modify nitrogen fluxes, soil depth and cattle pressure should influence ¹⁵N signatures. Our results suggest that perennial grasses are more conservative for nitrogen (inhibition of nitrification, less leaching via a perennial root system, slower cycling). The increase in leaf δ¹⁵N with N concentration is steeper in Loudetia togoensis than in the three other grasses. No significant difference was found between the ¹⁵N signatures of the four plots. Our results on ¹⁵N signatures and the fact that perennial grasses are much more abundant in the plots that are less grazed and have deeper soils, confirm that the switch from perennial to annual grasses is linked to a degradation in soil fertility and pasture quality. This suggests that ¹⁵N signatures can be used as indicators of fertility.     

Foliar and soil δ15N values reveal increased nitrogen partitioning among species in diverse grassland communities

Plant and soil nitrogen isotope ratios (δ¹⁵N) were studied in experimental grassland plots of varying species richness. We hypothesized that partitioning of different sources of soil nitrogen among four plant functional groups (legumes, grasses, small herbs, tall herbs) should increase with diversity. Four years after sowing, all soils were depleted in ¹⁵N in the top 5 cm whereas in non‐legume plots soils were enriched in ¹⁵N at 5–25 cm depth. Decreasing foliar δ¹⁵N and Δδ¹⁵N (= foliar δ¹⁵N ? soil δ¹⁵N) values in legumes indicated increasing symbiotic N₂ fixation with increasing diversity. In grasses, foliar Δδ¹⁵N also decreased with increasing diversity suggesting enhanced uptake of N depleted in ¹⁵N. Foliar Δδ¹⁵N values of small and tall herbs were unaffected by diversity. Foliar Δδ¹⁵N values of grasses were also reduced in plots containing legumes, indicating direct use of legume‐derived N depleted in ¹⁵N. Increased foliar N concentrations of tall and small herbs in plots containing legumes without reduced foliar δ¹⁵N indicated that these species obtained additional mineral soil N that was not consumed by legumes. These functional group and species specific shifts in the uptake of different N sources with increasing diversity indicate complementary resource use in diverse communities.