δ15N of forest soil and understorey vegetation reflect the former agricultural land use

Since the middle of the 19th century, the area covered by forests in France has doubled. These new forests grow on previous agricultural lands. We have studied the influence of this agricultural history on the ¹⁵N abundance of present-day forests planted on farmlands in the Vosges mountains (north-eastern France) between 1898 and 1930. Different types of land use were identified from old cadastres (1814–1836) of 16 farms. Ancient forests adjacent to farmlands were used as controls. Former pastures, meadows, croplands, gardens and ancient forests were compared for soil δ¹⁵N (fraction <50 μm and total soil), C/N, P and N content and fern (Dryopteris carthusiana) δ¹⁵N. The mean δ¹⁵N of soil increased in the order ancient forests (+0.0‰)<pastures (+1.4‰)<croplands (+1.6‰)<meadows (+2.5‰)<gardens (+3.8‰). This increase in soil δ¹⁵N with the intensity of former land use was related to the former input of ¹⁵N-enriched manure, and to an activation of soil nitrification leading to ¹⁵N-depleted nitrate export on previously manured parcels. Fern δ¹⁵N increased in the same order as soil δ¹⁵N in relation to past land use. The mean δ¹⁵N of fern in ancient forests (–4.4‰) and former pastures (–3.4‰) was 5‰ lower than soil δ¹⁵N and the two variables were strongly correlated. The δ¹⁵N of fern in formerly manured parcels varied little (cropland: –2.7‰, meadows: –2.6‰ and gardens: –2.2‰) and independently of soil δ¹⁵N, suggesting that the soil sources of fern N differed between unmanured and manured parcels. Understorey plant δ¹⁵N and soil δ¹⁵N appear to be excellent tracers of previous land use in forests, and could be used in historical studies. The persistence of high isotopic ratios in previously manured parcels, almost a century after afforestation, suggests a long-term influence of former land use on the N cycle in forest soils. Received: 22 January 1999 / Accepted: 22 July 1999     

Hydrogen and oxygen isotope ratios of tree ring cellulose for field-grown riparian trees

The isotopic composition of tree ring cellulose was obtained over a 2-year period from small-diameter riparian-zone trees at field sites that differed in source water isotopic composition and humidity. The sites were located in Utah (cool and low humidity), Oregon (cool and high humidity), and Arizona (warm and low humidity) with source water isotope ratio values of –125/–15‰ (δD/δ¹⁸O), –48/–6‰, and –67/–7‰, respectively. Monthly environmental measurements included temperature and humidity along with measurements of the isotope ratios in atmospheric water vapor, stream, stem, and leaf water. Small riparian trees used only stream water (both δD and δ¹⁸O of stem and stream water did not differ), but δ values of both atmospheric water vapor and leaf water varied substantially between months. Differences in ambient temperature and humidity conditions between sites contributed to substantial differences in leaf water evaporative enrichment. These leaf water differences resulted in differences in the δD and δ¹⁸O values of tree ring cellulose, indicating that humidity information was recorded in the annual rings of trees. These environmental and isotopic measurements were used to test a mechanistic model of the factors contributing to δD and δ¹⁸O values in tree ring cellulose. The model was tested in two parts: (a) a leaf water model using environmental information to predict leaf water evaporative enrichment and (b) a model describing biochemical fractionation events and isotopic exchange with medium water. The models adequately accounted for field observations of both leaf water and tree ring cellulose, indicating that the model parameterization from controlled experiments was robust even under uncontrolled and variable field conditions. Received: 7 April 1999 / Accepted: 8 December 1999     

Nitrogen and carbon isotope responses of Chinese cabbage and chrysanthemum to the application of liquid pig manure

The effects of the liquid pig manure (LM) used in organic farming on the natural abundance of ¹⁵N and ¹³C signatures in plant tissues have not been studied. We hypothesized that application of LM will (1) increase δ¹⁵N of plant tissues due to the high δ¹⁵N of N in LM as compared with soil N or inorganic fertilizer N, and (2) increase δ¹³C of plant tissues as a result of high salt concentration in LM that decreases stomatal conductance of plants. To test these hypotheses, variations in the δ¹⁵N and δ¹³C of Chinese cabbage (Brassica campestris L.) and chrysanthemum (Chrysanthemum morifolium Ramatuelle) with two different LMs (with δ¹⁵N of +15.6 and +18.2‰) applied at two rates (323 and 646 kg N ha^-1 for cabbage and 150 and 300 kg N ha^-1 for chrysanthemum), or urea (δ¹⁵N = -2.7‰) applied at the lower rate above for the respective species, in addition to the control (no N input) were investigated through a 60-day pot experiment. Application of LM significantly increased plant tissue δ¹⁵N (range +9.4 to +14.9‰) over the urea (+3.2 to +3.3‰) or control (+6.8 to 7.7‰) treatments regardless of plant species, strongly reflecting the δ¹⁵N of the N source. Plant tissue δ¹³C were not affected by the treatments for cabbage (range −30.8 to −30.2‰) or chrysanthemum (−27.3 to −26.8‰). However, cabbage dry matter production decreased while its δ¹³C increased with increasing rate of LM application or increasing soil salinity (P < 0.05), suggesting that salinity stress caused by high rate of LM application likely decreased stomatal conductance and limited growth of cabbage. Our study expanded the use of the δ¹⁵N technique in N source (organic vs. synthetic fertilizer) identification and suggested that plant tissue δ¹³C maybe a sensitive indicator of plant response to salinity stress caused by high LM application rates.     

Interseasonal comparison of CO2 concentrations, isotopic composition, and carbon dynamics in an Amazonian rainforest (French Guiana)

Canopy CO₂ concentrations in a tropical rainforest in French Guiana were measured continuously for 5 days during the 1994 dry season and the 1995 wet season. Carbon dioxide concentrations ([CO₂]) throughout the canopy (0.02–38 m) showed a distinct daily pattern, were well-stratified and decreased with increasing height into the canopy. During both seasons, daytime [CO₂] in the upper and middle canopy decreased on average 7–10 μmol mol⁻¹ below tropospheric baseline values measured at Barbados. Within the main part of the canopy (≥ 0.7 m), [CO₂] did not differ between the wet and dry seasons. In contrast, [CO₂] below 0.7 m were generally higher during the dry season, resulting in larger [CO₂] gradients. Supporting this observation, soil CO₂ efflux was on average higher during the dry season than during the wet season, either due to diffusive limitations and/or to oxygen deficiency of root and microbial respiration. Soil respiration rates decreased by 40% after strong rain events, resulting in a rapid decrease in canopy [CO₂] immediately above the forest floor of about 50␣μmol mol⁻¹. Temporal and spatial variations in [CO₂]_canopy were reflected in changes of δ¹³C_canopy and δ¹⁸O_canopy values. Tight relationships were observed between δ¹³C and δ¹⁸O of canopy CO₂ during both seasons (r ² > 0.86). The most depleted δ¹³C_canopy and δ¹⁸O_canopy values were measured immediately above the forest floor (δ¹³C = −16.4‰; δ¹⁸O = 39.1‰ SMOW). Gradients in the isotope ratios of CO₂ between the top of the canopy and the forest floor ranged between 2.0‰ and 6.3‰ for δ¹³C, and between 1.0‰ and 3.5‰ for δ¹⁸O. The δ¹³C_leaf and calculated c _i/c _a of foliage at three different positions were similar for the dry and wet seasons indicating that the canopy maintained a constant ratio of photosynthesis to stomatal conductance. About 20% of the differences in δ¹³C_leaf within the canopy was accounted for by source air effects, the remaining 80% must be due to changes in c _i/c _a. Plotting 1/[CO₂] vs. the corresponding δ¹³C ratios resulted in very tight, linear relationships (r ² = 0.99), with no significant differences between the two seasons, suggesting negligible seasonal variability in turbulent mixing relative to ecosystem gas exchange. The intercepts of these relationships that should be indicative of the δ¹³C of respired sources were close to the measured δ¹³C of soil respired CO₂ and to the δ¹³C of litter and soil organic matter. Estimates of carbon isotope discrimination of the entire ecosystem, Δ_e, were calculated as 20.3‰ during the dry season and as 20.5‰ during the wet season. Received: 3 March 1996 / Accepted: 19 October 1996     

Influence of stand structure on carbon-13 of vegetation, soils, and canopy air within deciduous and evergreen forests in Utah, United States

Carbon isotope ratios (δ¹³C) were studied in evergreen and deciduous forest ecosystems in semi-arid Utah (Pinus contorta, Populus tremuloides, Acer negundo and Acer grandidentatum). Measurements were taken in four to five stands of each forest ecosystem differing in overstory leaf area index (LAI) during two consecutive growing seasons. The δ¹³C_leaf (and carbon isotope discrimination) of understory vegetation in the evergreen stands (LAI 1.5–2.2) did not differ among canopies with increasing LAI, whereas understory in the deciduous stands (LAI 1.5–4.5) exhibited strongly decreasing δ¹³C_leaf values (increasing carbon isotope discrimination) with increasing LAI. The δ¹³C values of needles and leaves at the top of the canopy were relatively constant over the entire LAI range, indicating no change in intrinsic water-use efficiency with overstory LAI. In all canopies, δ¹³C_leaf decreased with decreasing height above the forest floor, primarily due to physiological changes affecting c _i/c _a (> 60%) and to a minor extent due to δ¹³C of canopy air (< 40%). This intra-canopy depletion of δ¹³C_leaf was lowest in the open stand (1‰) and greatest in the denser stands (4.5‰). Although overstory δ¹³C_leaf did not change with canopy LAI, δ¹³C of soil organic carbon increased with increasing LAI in Pinus contorta and Populus tremuloides ecosystems. In addition, δ¹³C of decomposing organic carbon became increasingly enriched over time (by 1.7–2.9‰) for all deciduous and evergreen dry temperate forests. The δ¹³C_canopy of CO₂ in canopy air varied temporally and spatially in all forest stands. Vertical canopy gradients of δ¹³C_canopy, and [CO₂]_canopy were larger in the deciduous Populus tremuloides than in the evergreen Pinu contorta stands of similar LAI. In a very wet and cool year, ecosystem discrimination (Δ_e) was similar for both deciduous Populus tremulodies (18.0 ± 0.7‰) and evergreen Pinus contorta (18.3 ± 0.9‰) stands. Gradients of δ¹³C_canopy and [CO₂]_canopy were larger in denser Acer spp. stands than those in the open stand. However, ¹³C enrichment above and photosynthetic draw-down of [CO₂]_canopy below tropospheric baseline values were larger in the open than in the dense stands, due to the presence of a vigorous understory vegetation. Seasonal patterns of the relationship δ¹³C_canopy versus 1/[CO₂]_canopy were strongly influenced by precipitation and air temperature during the growing season. Estimates of Δ_e for Acer spp. did not show a significant effect of stand structure, and averaged 16.8 ± 0.5‰ in 1933 and 17.4 ± 0.7‰ in 1994. However, Δ_e varied seasonally with small fluctuations for the open stand (2‰), but more pronounced changes for the dense stand (5‰). Received: 15 April 1996 / Accepted: 19 October 1996     

Variations in leaf β13C along a vertical profile of irradiance in a temperate Japanese forest

The vertical profile of stable carbon isotope ratios (δ¹³C) of leaves was analyzed for 13 tree species in a cool-temperate deciduous forest in Japan. The vertical distribution of long-term averaged δ¹³C in atmospheric CO₂ (δ_a) was estimated from δ¹³C of dry matter from NADP-malic enzyme type C₄ plant (Zea mays L. var. saccharata Sturt.) grown at a tower in the forest for 32␣days, assuming constant Δ value (3.3‰) in Z. mays against height. The δ_a value obtained from δ¹³C in Z.␣mays was lowest at the forest floor (−9.30 ± 0.03‰), increased with height, and was almost constant above 10␣m (−7.14 ± 0.14‰). Then leaf Δ values for the tree species were calculated from tree leaf δ¹³ C andδ_a. Mean leaf Δ values for the three tall deciduous species (Fraxinus mandshurica, Ulmus davidiana, and Alnus hirsuta) were significantly different among three height levels in the forest: 23.1 ± 0.7‰ at the forest floor (understory), 21.4 ± 0.5‰ in lower canopy, and 20.5 ± 0.3‰ in upper canopy. The true difference in tree leaf Δ among the forest height levels might be even greater, because Δ in Z. mays probably increased with shading by up to ∼‰. The difference in tree leaf Δ among the forest height levels would be mainly due to decreasing intercellular CO₂ (C _i) with the increase in irradiance. Potential assimilation rate for the three tree species probably increased with height, since leaf nitrogen content on an area basis for these species also increased with height. However, the increase in stomatal conductance for these tree species would fail to meet the increase in potential assimilation rate, which might lead to increasing the degree of stomatal limitation in photosynthesis with height. Received: 30 September 1995 / Accepted: 25 October 1996     

Effects of climate, tree age, dominance and growth on δ15N in young pinewoods

Needles, annual rings from basal stem discs and bark of three dominant and three suppressed Pinus pinaster from a 12-year-old pine stand (naturally regenerated after a wildfire) were analysed to study the effects of climate, tree age, dominance, and growth on tree δ¹⁵N. Foliar-N concentration in dominant pines (0.780–1.474% N) suggested that soil N availability was sufficient, a circumstance that allowed isotopic discrimination by plants and (greater) differences in δ¹⁵N among trees. The δ¹⁵N decreases in the order wood (−0.20 to +6.12‰), bark (−1.84 to +1.85‰) and needles (−2.13 to +0.77‰). In all trees, before dominance establishment (years 1–8), the N stored in each ring displayed a decreasing δ¹⁵N tendency as the tree grows, which is mainly due to a more “closed” N cycle or an increasing importance of N sources with lower δ¹⁵N. After dominance establishment (years 9–12), wood δ¹⁵N values were higher in suppressed than in dominant trees (2.62 and 1.46‰, respectively; P < 0.01) while the reverse was true for needles and bark; simultaneously, the absolute amount of N stored by suppressed pines in successive rings decreased, suggesting a lower soil N assimilation. These results could be explained by lignification acting as major N source for needles in suppressed pines because products released and reallocated during lignification are ¹⁵N-depleted compared with the source. According to principal component analysis, wood δ¹⁵N appears associated with wood N concentration and precipitation during the growing season, but clearly opposed to age, basal area increment and mean temperature in spring and summer.     

The carbon origin of arbuscular mycorrhizal fungi estimated from δ13C values of individual spores

 The origin of carbon in the spores of arbuscular mycorrhizal (AM) fungi was quantified based on their obligate symbiosis with C₃ and C₄ plants showing clearly different δ¹³C values. The δ¹³C values of individual spores of the AM fungus Gigaspora margarita were analyzed. In monoculture pots of a C₃ or a C₄ plant species, spore δ¹³C values were ca. 3.5‰ lower than those of host roots. In coculture pots of a C₃ and a C₄ plant species, spore δ¹³C values varied between those of the roots of C₃ and C₄ plants, and increased linearly from the C₃ to the proximity of the C₄ plant (P<0.01). This reflects the higher δ¹³C values in C₄ plants than in C₃ plants. Thus the carbon origin of G. margarita spores changed with growth state and combination of host plants. In the presence of fresh plant residue instead of living host plants, spore δ¹³C values did not vary with distance from the residue. This finding supports the current view that AM fungi are obligate symbionts. Accepted: 12 February 1999     

Correlations between foliar δ15N and nitrogen concentrations may indicate plant-mycorrhizal interactions

Nitrogen isotope measurements may provide insights into changing interactions among plants, mycorrhizal fungi, and soil processes across environmental gradients. Here, we report changes in δ¹⁵N signatures due to shifts in species composition and nitrogen (N) dynamics. These changes were assessed by measuring fine root biomass, net N mineralization, and N concentrations and δ¹⁵N of foliage, fine roots, soil, and mineral N across six sites representing different post-deglaciation ages at Glacier Bay, Alaska. Foliar δ¹⁵N varied widely, between 0 and –2‰ for nitrogen-fixing species, between 0 and –7‰ for deciduous non-fixing species, and between 0 and –11‰ for coniferous species. Relatively constant δ¹⁵N values for ammonium and generally low levels of soil nitrate suggested that differences in ammonium or nitrate use were not important influences on plant δ¹⁵N differences among species at individual sites. In fact, the largest variation among plant δ¹⁵N values were observed at the youngest and oldest sites, where soil nitrate concentrations were low. Low mineral N concentrations and low N mineralization at these sites indicated low N availability. The most plausible mechanism to explain low δ¹⁵N values in plant foliage was a large isotopic fractionation during transfer of nitrogen from mycorrhizal fungi to plants. Except for N-fixing plants, the foliar δ¹⁵N signatures of individual species were generally lower at sites of low N availability, suggesting either an increased fraction of N obtained from mycorrhizal uptake (f), or a reduced proportion of mycorrhizal N transferred to vegetation (T _r). Foliar and fine root nitrogen concentrations were also lower at these sites. Foliar N concentrations were significantly correlated with δ¹⁵N in foliage of Populus, Salix, Picea, and Tsuga heterophylla, and also in fine roots. The correlation between δ¹⁵N and N concentration may reflect strong underlying relationships among N availability, the relative allocation of carbon to mycorrhizal fungi, and shifts in either f or T _r. Received: 14 December 1998 / Accepted: 16 August 1999     

Effect of host shoot clipping on carbon and nitrogen sources for arbuscular mycorrhizal fungi

The effect of clipping of the host-plant shoot on the sources of carbon and nitrogen for the arbuscular mycorrhizal (AM) fungus Gigaspora margarita was determined by measuring ¹³C in spores and hyphae in cocultures of C₃ and C₄ plants and by differential ¹⁵N labeling. C₃ and C₄ plants, which have different δ¹³C values, were grown in the same container separated by a series of hyphal compartments. The C₃ and C₄ plants were applied with ¹⁴N- and ¹⁵N-urea, respectively. After clipping of the C₃ shoots, spore δ ¹³C gradually approached that of the C₄ roots. Hyphal δ ¹³C paralleled that of spores. Spore % ¹⁵N was similar to that of mineral N in the C₄ plant compartment. Thus C in G. margarita coming from the clipped plants decreased with time. This demonstrates that C in AM fungi comes from living plants, whilst the N in spores comes mostly from the soil. Accepted: 28 November 2000     

Isotopic signature of nitrate in two contrasting watersheds of Brush Brook,Vermont, USA

We used the dual isotope method to study differences in nitrate export in two subwatersheds in Vermont, USA. Precipitation, soil water and streamwater samples were collected from two watersheds in Camels Hump State Forest, located within the Green Mountains of Vermont. These samples were analyzed for the δ¹⁵N and δ¹⁸O of NO₃⁻. The range of δ¹⁵N–NO₃⁻ values overlapped, with precipitation −4.5‰ to +2.0‰ (n = 14), soil solution −10.3‰ to +6.2‰ (n = 12) and streamwater +0.3‰ to +3.1‰ (n = 69). The δ¹⁸O of precipitation NO₃⁻ (mean 46.8 ± 11.5‰) was significantly different (P < 0.001) from that of the stream (mean 13.2 ± 4.3‰) and soil waters (mean 14.5 ± 4.2‰) even during snowmelt periods. Extracted soil solution and streamwater δ¹⁸O of NO₃⁻ were similar and within the established range of microbially produced NO₃⁻, demonstrating that NO₃⁻ was formed by microbial processes. The δ¹⁵N and δ¹⁸O of NO₃⁻ suggests that although the two tributaries have different seasonal NO₃⁻ concentrations, they have a similar NO₃⁻ source.     

Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients

 Our objective was to evaluate the relative importance of gradients in light intensity and the isotopic composition of atmospheric CO₂ for variation in leaf carbon isotope ratios within a Pinus resinosa forest. In addition, we measured photosynthetic gas exchange and leaf carbon isotope ratios on four understory species (Dryopteris carthusiana, Epipactus helleborine, Hieracium floribundum, Rhamnus frangula), in order to estimate the consequence of the variation in the understory light microclimate for carbon gain in these plants. During midday, CO₂ concentration was relatively constant at vertical positions ranging from 15 m to 3 m above ground. Only at positions below 3 m was CO₂ concentration significantly elevated above that measured at 15 m. Based on the strong linear relationship between changes in CO₂ concentration and δ¹³C values for air samples collected during a diurnal cycle, we calculated the expected vertical profile for the carbon isotope ratio of atmospheric CO₂ within the forest. These calculations indicated that leaves at 3 m height and above were exposed to CO₂ of approximately the same isotopic composition during daylight periods. There was no significant difference between the daily mean δ¹³C values at 15 m (–7.77‰) and 3 m (–7.89‰), but atmospheric CO₂ was significantly depleted in ¹³C closer to the ground surface, with daily average δ¹³C values of –8.85‰ at 5 cm above ground. The light intensity gradient in the forest was substantial, with average photosynthetically active radiation (PAR) on the forest floor approximately 6% of that received at the top of the canopy. In contrast, there were only minor changes in air temperature, and so it is likely that the leaf-air vapour pressure difference was relatively constant from the top of the canopy to the forest floor. For red pine and elm tree samples, there was a significant correlation between leaf δ¹³C value and the height at which the leaf sample was collected. Leaf tissue sampled near the forest floor, on average, had lower δ¹³C values than samples collected near the top of the canopy. We suggest that the average light intensity gradient through the canopy was the major factor influencing vertical changes in tree leaf δ¹³C values. In addition, there was a wide range of variation (greater than 4‰) among the four understory plant species for average leaf δ¹³C values. Measurements of leaf gas exchange, under natural light conditions and with supplemental light, were used to estimate the influence of the light microclimate on the observed variation in leaf carbon isotope ratios in the understory plants. Our data suggest that one species, Epipactus helleborine, gained a substantial fraction of carbon during sunflecks. Received: 21 March 1996 / Accepted: 13 August 1996     

Effect of C3–C4 Vegetation Change on δ13C and δ15N Values of Soil Organic Matter Fractions Separated by Thermal Stability

Carbon isotopic composition of soils subjected to C₃–C₄ vegetation change can be used to estimate C turnover in bulk soil and in soil organic matter (SOM) pools with fast and intermediate turnover rates. We hypothesized that the biological availability of SOM pools is inversely proportional to their thermal stability, so that thermogravimetry can be used to separate SOM pools with contrasting turnover rates. Soil samples from a field plot cultivated for 10.5 years with the perennial C₄ plant Miscanthus×gigantheus were analyzed by thermogravimetry coupled with differential scanning calorimetry (DSC). Three SOM fractions were distinguished according to the differential weight losses and exothermic or endothermic reactions measured by DSC. The δ¹³C and δ¹⁵N values of these three fractions obtained by gradual soil heating were measured by IRMS. The weight losses up to 190 °C mainly reflected water evaporation because no significant C and N losses were detected and δ¹³C and δ¹⁵N values of the residual SOM remained unchanged. The δ¹³C values (−16.4‰) of SOM fraction decomposed between 190 and 390 °C (containing 79% of total soil C) were slightly closer to that of the Miscanthus plant tissues (δ¹³C = −11.8‰) compared to the δ¹³C values (−16.8‰) of SOM fraction decomposed above 390 °C containing the residual 21% of SOM. Thus, the C turnover in the thermally labile fraction was faster than that in thermally stable fractions, but the differences were not very strong. Therefore, in this first study combining TG-DSC with isotopic analysis, we conclude that the thermal stability of SOM was not very strongly related to biological availability of SOM fractions. In contrast to δ¹³C, the δ¹⁵N values strongly differed between SOM fractions, suggesting that N turnover in the soil was different from C turnover. More detailed fractionation of SOM by thermal analysis with subsequent isotopic analysis may improve the resolution for δ¹³C.     

Hydraulic Lift in Cork Oak Trees in a Savannah-Type Mediterranean Ecosystem and its Contribution to the Local Water Balance

The aim of this study was to identify the sources and depth of water uptake by 15-years old Quercus suber L. trees in southern Portugal under a Mediterranean climate, measuring δ¹⁸O and δD in the soil–plant-atmosphere continuum. Evidence for hydraulic lift was substantiated by the daily fluctuations observed in Ψ_s at 0.4 and 1 m depth and supported by similar δ¹⁸O values found in tree xylem sap, soil water in the rhizosphere and groundwater. From 0.25 m down to a depth of 1 m, δD trends differed according to vegetation type, showing a more depleted value in soil water collected under the evergreen trees (−47‰) than under dead grasses (−35‰). The hypothesis of a fractionation process occurring in the soil due to diffusion of water vapour in the dry soil is proposed to explain the more depleted soil δD signature observed under trees. Hydraulically lifted water was estimated to account for 17–81% of the water used during the following day by tree transpiration at the peak of the drought season, i.e., 0.1–14 L tree⁻¹ day⁻¹. Significant relationships found between xylem sap isotopic composition and leaf water potential in early September emphasized the positive impact of the redistribution of groundwater in the rhizosphere on tree water status.     

Dietary histories of herbivorous loricariid catfishes: evidence from δ<Superscript>13</Superscript>C values of otoliths

The ecology of many Neotropical fishes is difficult or often impossible to study during rainy seasons. Thus, ecological studies of tropical fishes are usually performed on fish captured only during dry seasons. Because otoliths preserve a record of life history, this study evaluated the utility of otolith stable isotope values for the investigation of trophic ecology of Neotropical fishes (specifically herbivorous loricariid catfish) throughout their lives. Because plant dietary materials have δ¹³C values that are determined by their photosynthetic pathways, metabolism and environmental conditions, different plants may impart different isotope values on fish otoliths that reflect consumption of these plants. The δ¹³C_(otolith) values of xylophagous Panaque nigrolineatus captured in the field were significantly lower than those of algivorous Hypostomus regani from a nearby region. A laboratory experiment wherein Hypostomus sp. had δ¹³C_(otolith) values that reflected the δ¹³C values of their plant diet and additional evidence indicate that δ¹³C_(otolith) values in loricariid catfish otoliths can record dietary history.     

Vascular plant 15N natural abundance in heath and forest tundra ecosystems is closely correlated with presence and type of mycorrhizal fungi in roots

In this study we show that the natural abundance of the nitrogen isotope 15, δ¹⁵N, of plants in heath tundra and at the tundra-forest ecocline is closely correlated with the presence and type of mycorrhizal association in the plant roots. A total of 56 vascular plant species, 7 moss species, 2 lichens and 6 species of fungi from four heath and forest tundra sites in Greenland, Siberia and Sweden were analysed for δ¹⁵N and N concentration. Roots of vascular plants were examined for mycorrhizal colonization, and the soil organic matter was analysed for δ¹⁵N, N concentration and soil inorganic, dissolved organic and microbial N. No arbuscular mycorrhizal (AM) colonizations were found although potential host plants were present in all sites. The dominant species were either ectomycorrhizal (ECM) or ericoid mycorrhizal (ERI). The δ¹⁵N of ECM or ERI plants was 3.5–7.7‰ lower than that of non-mycorrhizal (NON) species in three of the four sites. This corresponds to the results in our earlier study of mycorrhiza and plant δ¹⁵N which was limited to one heath and one fellfield in N Sweden. Hence, our data suggest that the δ¹⁵N pattern: NON/AM plants > ECM plants ≥ ERI plants is a general phenomenon in ecosystems with nutrient-deficient organogenic soils. In the fourth site, a␣birch forest with a lush herb/shrub understorey, the differences between functional groups were considerably smaller, and only the ERI species differed (by 1.1‰) from the NON species. Plants of all functional groups from this site had nearly twice the leaf N concentration as that found in the same species at the other three sites. It is likely that low inorganic N availability is a prerequisite for strong δ¹⁵N separation among functional groups. Both ECM roots and fruitbodies were ¹⁵N enriched compared to leaves which suggests that the difference in δ¹⁵N between plants with different kinds of mycorrhiza could be due to isotopic fractionation at the␣fungal-plant interface. However, differences in δ¹⁵N between soil N forms absorbed by the plants could also contribute to the wide differences in plant δ¹⁵N found in most heath and forest tundra ecosystems. We hypothesize that during microbial immobilization of soil ammonium the microbial N pool could become ¹⁵N-depleted and the remaining, plant-available soil ammonium ¹⁵N-enriched. The latter could be a main source of N for NON/AM plants which usually have high δ¹⁵N. In contrast, amino acids and other soil organic N compounds presumably are ¹⁵N-depleted, similar to plant litter, and ECM and ERI plants with high uptake of these N forms hence have low leaf δ¹⁵N. Further indications come from the δ¹⁵N of mosses and lichens which was similar to that of ECM plants. Tundra cryptogams (and ECM and ERI plants) have previously been shown to have higher uptake of amino acid than ammonium N; their low δ¹⁵N might therefore reflect the δ¹⁵N of free amino acids in the soil. The concentration of dissolved organic N was 3–16 times higher than that of inorganic N in the sites. Organic nitrogen could be an important N source for ECM and, in particular, ERI plants in heath and forest tundra ecosystems with low release rate of inorganic N from the soil organic matter. Received: 8 June 1997 / Accepted: 28 February 1998     

Relationships of stable carbon isotopes, plant water potential and growth: an approach to asses water use efficiency and growth strategies of dry land agroforestry species

The relationships between annual wood stable carbon isotope composition (δ¹³C), dry season midday plant water potential, and annual growth rate were investigated to asses the ability of agroforestry species to adapt to climate changes. 6–8 stem disks from four co-occurring species (Acacia senegal, A. seyal, A. tortilis and Balanites aegyptiaca) were collected for radial growth measurements using tree-ring analysis spanning 1930–2003. Annual δ¹³C was measured on three tree disks per species for the period 1970–2002. Midday plant water potential was measured during the dry season. Annual radial growth and midday plant water potential ranged from 0.27 to 9.12 mm and −1.0 to −5.0 MPa, respectively, with statistically significant differences. After correcting annual wood δ¹³C for atmospheric changes in δ¹³C, carbon isotopic composition ranged from −22.22 to −26.58‰. Relationships between δ¹³C, radial growth and plant water potentials revealed the interaction of water availability, stomatal conductance, δ¹³C values and growth. Two contrasting water use strategies and competitive advantages can be distinguished. Species with lower mean δ¹³C values (A. senegal and A. seyal) show high plant water potential and, hence, better growth during moist years. Thus, they indicate low water use efficiency (WUE) and opportunistic water use strategy. On the other hand, species with lower water potentials (A. tortilis and B. aegyptiaca) showed relative better growth performance and less increase in δ¹³C in drought years, reflecting their high WUE and conservative water use strategy. These results suggest that δ¹³C in tree rings can be useful in estimating historic changes in plant WUE and hence in screening drought tolerant species in the face of expected climate changes, as well as for assessing the functional diversity and risk reduction in mixed vegetation.     

Isotopic signatures and nutrient relations of plants inhabiting brackish wetlands in the northeastern coastal plain of Venezuela

The semi-diurnal tidal regime (≥2 m) in the Paria Gulf on the Atlantic coast of Venezuela, and the flat landscape of the region, allow the penetration for tens of km of marine waters into the rivers draining the northeastern coastal plain of the country. The levels of salinity, tidal flooding, and sedimentation decrease perpendicularly from the river channel toward the back swamps. The vegetation varies sequentially from fringe mangroves along the river margins, to back swamps containing forests dominated by Pterocarpus officinalis, herbaceous communities of Lagenocarpus guianensis, and palm swamps with Mauritia flexuosa, Chrysobalanus icaco, and Tabebuia spp. This environmental structure was used to test the hypotheses that: (a) mangrove distribution is strongly associated with salinity of interstitial water, and (b) they occupy areas where tidal influence and sediment dynamics determine a relatively open N cycle. Analyses of soil, water, and plants along a 1.5 km transect located near the confluence of the Guanoco and San Juan Rivers (Sucre and Monagas States, Venezuela) revealed that: (a) conductivity decreased from 11 to 0.2 mmhos cm⁻¹ from the river fringe to the internal swamp, whereas Na in the same stretch decreased from 100 to 2 μM; (b) average leaf tissue concentrations of Na, P, and N decreased significantly along the transect; (c) P. officinalis showed a large Na-exclusion capacity indicated by positive K/Na ratios from 8 to 200, and Crinum erubescens counteracted Na by accumulating K above 1,000 mmol kg⁻¹; (d) leaves varied widely in δ ¹³C (−25.5 to −32‰) and δ ¹⁵N (4 to −10.5‰) values. Samples were aggregated according to soil carbon content corresponding to those of the mangrove forest belt (5–28 mol C kg⁻¹; 0–650 from river fringe) and those of the back swamps (40–44 mol C kg⁻¹; 700–1,500 m from river fringe). The concentrations of Na, P, and N (in mmol kg⁻¹) and δ ¹⁵N values (in ‰) were significantly higher in the mangrove forest compared to the back swamp (Na 213 vs. 88; P 41 vs. 16; N 1,535 vs. 727; δ ¹⁵N 1.5 vs. −3.7), indicating that the fringe forest was not nutrient limited. These results support the hypotheses that mangroves are restricted to the more-saline sections of the transect, and that the fringe forest has a more open N cycle, favoring ¹⁵N accumulation within the system.     

Variation in woody plant δ13C along a topoedaphic gradient in a subtropical savanna parkland

δ¹³C values of C₃ plants are indicators of plant carbon–water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant δ¹³C at the landscape scale. We determined variation in leaf δ¹³C, leaf nitrogen per leaf area (N_area), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 × 12-m belt transect spanning an upland–lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The δ¹³C values of Prosopis glandulosa (deciduous N₂-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1–4‰ with decreasing elevation, with the δ¹³C value of P. glandulosa leaves being 1–3‰ higher than those of the two shrub species. Contrary to theory and results from previous studies, δ¹³C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N_area was positively correlated with leaf δ¹³C of all species (p < 0.01) and appeared to exert the strongest control over δ¹³C along this topoedaphic gradient. Since leaf N_area is positively related to photosynthetic capacity, plants with high leaf N_area are likely to have low p _I/p _a ratios and therefore higher δ¹³C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf δ¹³C. Following a progressive growing season drought in July/August, leaf δ¹³C decreased. The lower δ¹³C in August may reflect the accumulation of ¹³C-depleted epicuticular leaf wax. We suggest control of leaf δ¹³C along this topoedaphic gradient is mediated by leaf N_area rather than by stomatal conductance limitations associated with water availability.     

Ecosystem N Distribution and δ<Superscript>15</Superscript>N during a Century of Forest Regrowth after Agricultural Abandonment

Abstract Stable isotope ratios of terrestrial ecosystem nitrogen (N) pools reflect internal processes and input–output balances. Disturbance generally increases N cycling and loss, yet few studies have examined ecosystem δ¹⁵N over a disturbance-recovery sequence. We used a chronosequence approach to examine N distribution and δ¹⁵N during forest regrowth after agricultural abandonment. Site ages ranged from 10 to 115 years, with similar soils, climate, land-use history, and overstory vegetation (white pine Pinus strobus). Foliar N and δ¹⁵N decreased as stands aged, consistent with a progressive tightening of the N cycle during forest regrowth on agricultural lands. Over time, foliar δ¹⁵N became more negative, indicating increased fractionation along the mineralization–mycorrhizal–plant uptake pathway. Total ecosystem N was constant across the chronosequence, but substantial internal N redistribution occurred from the mineral soil to plants and litter over 115 years (>25% of ecosystem N or 1,610 kg ha⁻¹). Temporal trends in soil δ¹⁵N generally reflected a redistribution of depleted N from the mineral soil to the developing O horizon. Although plants and soil δ¹⁵N are coupled over millennial time scales of ecosystem development, our observed divergence between plants and soil suggests that they can be uncoupled during the disturbance-regrowth sequence. The approximate 2‰ decrease in ecosystem δ¹⁵N over the century scale suggests significant incorporation of atmospheric N, which was not detected by traditional ecosystem N accounting. Consideration of temporal trends and disturbance legacies can improve our understanding of the influence of broader factors such as climate or N deposition on ecosystem N balances and δ¹⁵N. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.