Water relations of evergreen and drought-deciduous trees along a seasonally dry tropical forest chronosequence

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (Ψ_L) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although Ψ_L did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ¹⁸O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar ¹⁸O (∆¹⁸O_l) and ¹³C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.     

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.     

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.     

Exotic plant communities shift water-use timing in a shrub-steppe ecosystem

Semiarid areas in the US have realized extensive and persistent exotic plant invasions. Exotics may succeed in arid regions by extracting soil water at different times or from different depths than native plants, but little data is available to test this hypothesis. Using estimates of root mass, gravimetric soil water, soil-water potential, and stable isotope ratios in soil and plant tissues, we determined water-use patterns of exotic and native plant species in exotic- and native-dominated communities in Washington State, USA. Exotic and native communities both extracted 12 ± 2 cm of water from the top 120 cm of soil during the growing season. Exotic communities, however, shifted the timing of water use by extracting surface (0–15 cm) soil water early in the growing season (i.e., April to May) before native plants were active, and by extracting deep (0–120 cm) soil water late in the growing season (i.e., June to July) after natives had undergone seasonal senescence. We found that δ ¹⁸O values of water in exotic annuals (e.g., −11.8 ± 0.4 ‰ for Bromus tectorum L.) were similar to δ ¹⁸O values of surface soil water (e.g., −13.3 ± 1.4 ‰ at −15 cm) suggesting that transpiration by these species explained early season, surface water use in exotic communities. We also found that δ ¹⁸O values of water in taprooted exotics (e.g., −17.4 ± 0.3 ‰ for Centaurea diffusa Lam.) were similar to δ ¹⁸O values of deep soil water (e.g., −18.4 ± 0.1 ‰ at −120 cm) suggesting that transpiration by these species explained late season, deep water use. The combination of early-season, shallow water-use by exotic winter-actives and late-season, deep water-use by taprooted perennials potentially explains how exotic communities resist establishment of native species that largely extracted soil water only in the middle of the growing season (i.e., May to June). Early season irrigation or the planting of natives with established root systems may allow native plant restoration.     

Seasonal plant water uptake patterns in the saline southeast Everglades ecotone

The purpose of this study was to determine the seasonal water use patterns of dominant macrophytes coexisting in the coastal Everglades ecotone. We measured the stable isotope signatures in plant xylem water of Rhizophora mangle, Cladium jamaicense, and Sesuvium portulacastrum during the dry (DS) and wet (WS) seasons in the estuarine ecotone along Taylor River in Everglades National Park, FL, USA. Shallow soilwater and deeper groundwater salinity was also measured to extrapolate the salinity encountered by plants at their rooting zone. Average soil water oxygen isotope ratios (δ ¹⁸O) was enriched (4.8 ± 0.2‰) in the DS relative to the WS (0.0 ± 0.1‰), but groundwater δ ¹⁸O remained constant between seasons (DS: 2.2 ± 0.4‰; WS: 2.1 ± 0.1‰). There was an inversion in interstitial salinity patterns across the soil profile between seasons. In the DS, shallow water was euhaline [i.e., 43 practical salinity units (PSU)] while groundwater was less saline (18 PSU). In the WS, however, shallow water was fresh (i.e., 0 PSU) but groundwater remained brackish (14 PSU). All plants utilized 100% (shallow) freshwater during the WS, but in the DS R. mangle switched to a soil–groundwater mix (δ 55% groundwater) while C. jamaicense and S. portulacastrum continued to use euhaline shallow water. In the DS, based on δ ¹⁸O data, the roots of R. mangle roots were exposed to salinities of 25.4 ± 1.4 PSU, less saline than either C. jamaicense (39.1 ± 2.2 PSU) or S. portulacastrum (38.6 ± 2.5 PSU). Although the salinity tolerance of C. jamaicense is not known, it is unlikely that long-term exposure to high salinity is conducive to the persistence of this freshwater marsh sedge. This study increases our ecological understanding of how water uptake patterns of individual plants can contribute to ecosystem levels changes, not only in the southeast saline Everglades, but also in estuaries in general in response to global sea level rise and human-induced changes in freshwater flows.     

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     

Carbon nutrition of mature green orchid <Emphasis Type="Italic">Serapias strictiflora</Emphasis> and its mycorrhizal fungus <Emphasis Type="Italic">Epulorhiza</Emphasis> sp.

We studied the nutritional modes of the orchid Serapias strictiflora and its mycorrhizal fungus Epulorhiza sp. using the differences in carbon isotopic composition (δ¹³C) of C₃ orchid and C₄ maize tissues. We found that if cultivated in substrate lacking any organic compounds, the mycorrhizal extraradical mycelia (δ¹³C = −26.3 ± 0.2 ‰) developed well, despite being fully dependent on nutrition from orchid roots (δ¹³C = −28.6 ± 0.1 ‰). If the mycorrhizal fungus had additional access to and colonized decaying maize roots (δ¹³C = −14.6 ± 0.1 ‰), its isotopic composition (δ¹³C = −21.6 ± 0.4 ‰) reflected a mixture of biotrophy and saprotrophy. No statistically significant differences in δ¹³C of new storage tubers were found between Epulorhiza-associated orchids with (δ¹³C = -28.2 ± 0.1 ‰) and without access to maize roots (δ¹³C = −28.6 ± 0.2 ‰). We conclude that autotrophy is the predominant nutritional mode of mature S. strictiflora plants and that they supply their mycorrhizal fungus with substantial amount of carbon (69 ± 3 % of the fungus demand), even if the fungus feeds saprotrophically.     

Natural 15N abundance of soil N pools and N2O reflect the nitrogen dynamics of forest soils

Natural ¹⁵N abundance measurements of ecosystem nitrogen (N) pools and ¹⁵N pool dilution assays of gross N transformation rates were applied to investigate the potential of δ¹⁵N signatures of soil N pools to reflect the dynamics in the forest soil N cycle. Intact soil cores were collected from pure spruce (Picea abies (L.) Karst.) and mixed spruce-beech (Fagus sylvatica L.) stands on stagnic gleysol in Austria. Soil δ¹⁵N values of both forest sites increased with depth to 50 cm, but then decreased below this zone. δ¹⁵N values of microbial biomass (mixed stand: 4.7 ± 0.8‰, spruce stand: 5.9 ± 0.9‰) and of dissolved organic N (DON; mixed stand: 5.3 ± 1.7‰, spruce stand: 2.6 ± 3.3‰) were not significantly different; these pools were most enriched in ¹⁵N of all soil N pools. Denitrification represented the main N₂O-producing process in the mixed forest stand as we detected a significant ¹⁵N enrichment of its substrate NO₃⁻ (3.6 ± 4.5‰) compared to NH₄⁺ (−4.6 ± 2.6‰) and its product N₂O (−11.8 ± 3.2‰). In a ¹⁵N-labelling experiment in the spruce stand, nitrification contributed more to N₂O production than denitrification. Moreover, in natural abundance measurements the NH₄⁺ pool was slightly ¹⁵N-enriched (−0.4 ± 2.0 ‰) compared to NO₃⁻ (−3.0 ± 0.6 ‰) and N₂O (−2.1 ± 1.1 ‰) in the spruce stand, indicating nitrification and denitrification operated in parallel to produce N₂O. The more positive δ¹⁵N values of N₂O in the spruce stand than in the mixed stand point to extensive microbial N₂O reduction in the spruce stand. Combining natural ¹⁵N abundance and ¹⁵N tracer experiments provided a more complete picture of soil N dynamics than possible with either measurement done separately.     

Partitioning of soil water among canopy trees in a seasonally dry tropical forest

Little is known about partitioning of soil water resources in species-rich, seasonally dry tropical forests. We assessed spatial and temporal patterns of soil water utilization in several canopy tree species on Barro Colorado Island, Panama, during the 1997 dry season. Stable hydrogen isotope composition (δD) of xylem and soil water, soil volumetric water content (θ_v), and sap flow were measured concurrently. Evaporative fractionation near the soil surface caused soil water δD to decrease from about –15‰ at 0.1 m to –50 to –55‰ at 1.2 m depth. Groundwater sampled at the sources of nearby springs during this period yielded an average δD value of –60‰. θ_v increased sharply and nearly linearly with depth to 0.7 m, then increased more slowly between 0.7 and 1.05 m. Based on xylem δD values, water uptake in some individual plants appeared to be restricted largely to the upper 20 cm of the soil profile where θ_v dropped below 20% during the dry season. In contrast, other individuals appeared to have access to water at depths greater than 1 m where θ_v remained above 45% throughout the dry season. The depths of water sources for trees with intermediate xylem δD values were less certain because variation in soil water δD between 20 and 70 cm was relatively small. Xylem water δD was also strongly dependent on tree size (diameter at breast height), with smaller trees appearing to preferentially tap deeper sources of soil water than larger trees. This relationship appeared to be species independent. Trees able to exploit progressively deeper sources of soil water during the dry season, as indicated by increasingly negative xylem δD values, were also able to maintain constant or even increase rates of water use. Seasonal courses of water use and soil water partitioning were associated with leaf phenology. Species with the smallest seasonal variability in leaf fall were also able to tap increasingly deep sources of soil water as the dry season progressed. Comparison of xylem, soil, and groundwater δD values thus pointed to spatial and temporal partitioning of water resources among several tropical forest canopy tree species during the dry season. Received: 5 October 1998 / Accepted: 23 June 1999     

Vegetation control effects on untreated wood,crude cellulose and holocellulose δ<Superscript>13</Superscript>C of early and latewood in 3- to 5-year-old rings of Douglas-fir

The stable carbon (C) composition of tree rings expressed as δ¹³C, is a measure of intrinsic water-use efficiency and can indicate the occurrence of past water shortages for tree growth. We examined δ¹³C in 3- to 5-year-old rings of Douglas-fir (Pseudotsuga menziesii (Mirb) Franco) trees to elucidate if decreased water supply or uptake was a critical factor in the observed growth reduction of trees competing with understory herb and shrub vegetation compared to those growing without competition. We hypothesized that there would be no differences in δ¹³C of earlywood in trees growing in plots with competing vegetation and those in plots receiving complete vegetation control during 5 years because earlywood formed early in the growing season when soil water was ample. We also hypothesized that δ¹³C in latewood which was formed during the later half of the growing season when precipitation was low, would be greater (less negative) in trees in plots without vegetation control. We then separated early and latewood from rings for three consecutive years and analyzed their δ¹³C composition. No significant differences in earlywood δ¹³C in years 3–5 were observed for trees in the two vegetation control treatments. δ¹³C of untreated latewood separated from wood cores was greater in 4- and 5-year-old rings of trees growing with competing vegetation compared to trees growing without vegetation competition (i.e., −25.5 vs. −26.3‰ for year 4, and −26.1 vs. −26.8‰ for year 5). Results suggest that water shortages occurred in Douglas-fir trees on this coastal Washington site in the latewood-forming portion of the growing season of years 4 and 5 in the no-vegetation control treatment. We also compared δ¹³C from untreated wood, crude cellulose extracted with the Diglyme–HCl method, and holocellulose extracted with toluene–ethanol to see if the extraction method would increase the sensitivity of the analysis. δ¹³C values from the two extraction methods were highly correlated with those from untreated samples (r ² = 0.97, 0.98, respectively). Therefore, using untreated wood would be as effective as using crude cellulose or holocellulose to investigate δ¹³C patterns in young Douglas-fir.     

An isotopic method for testing the influence of leaf litter quality on carbon fluxes during decomposition

During microbial breakdown of leaf litter a fraction of the C lost by the litter is not released to the atmosphere as CO₂ but remains in the soil as microbial byproducts. The amount of this fraction and the factors influencing its size are not yet clearly known. We performed a laboratory experiment to quantify the flow of C from decaying litter into the soil, by means of stable C isotopes, and tested its dependence on litter chemical properties. Three sets of ¹³C-depleted leaf litter (Liquidambar styraciflua L., Cercis canadensis L. and Pinus taeda L.) were incubated in the laboratory in jars containing ¹³C-enriched soil (i.e. formed C4 vegetation). Four jars containing soil only were used as a control. Litter chemical properties were measured using thermogravimetry (Tg) and pyrolysis–gas chromatography/mass spectrometry–combustion interface–isotope ratio mass spectrometry (Py–GC/MS–C–IRMS). The respiration rates and the δ¹³C of the respired CO₂ were measured at regular intervals. After 8 months of incubation, soils incubated with both L. styraciflua and C. canadensis showed a significant change in δ¹³C (δ¹³C_final = −20.2 ± 0.4‰ and −19.5 ± 0.5‰, respectively) with respect to the initial value (δ¹³C_initial = −17.7 ± 0.3‰); the same did not hold for soil incubated with P. taeda (δ¹³C_final:−18.1 ± 0.5‰). The percentages of litter-derived C in soil over the total C loss were not statistically different from one litter species to another. This suggests that there is no dependence of the percentage of C input into the soil (over the total C loss) on litter quality and that the fractional loss of leaf litter C is dependent only on the microbial assimilation efficiency. The percentage of litter-derived C in soil was estimated to be 13 ± 3% of total C loss.     

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.     

Carbon and nitrogen isotope composition of vegetation on King George Island,maritime Antarctic

We report abundance of ¹³C and ¹⁵N contents in terrestrial plants (mosses, lichens, liverworts, algae and grasses) from the area of Barton Peninsula (King George Island, maritime Antarctic). The investigated plants show a wide range of δ¹³C and δ¹⁵N values between −29.0 and −20.0‰ and between −15.3 and 22.8‰, respectively. The King George Island terrestrial plants show species specificity of both carbon and nitrogen isotope compositions, probably due to differences in plant physiology and biochemistry, related to their sources and in part to water availability. Carbon isotope compositions of Antarctic terrestrial plants are typical of the C₃ photosynthetic pathway. Lichens are characterized by the widest carbon isotope range, from −29.0 to −20.0‰. However, the average δ¹³C value of lichens is the highest (−23.6 ± 2.8‰) among King George Island plants, followed by grasses (−25.6 ± 1.7‰), mosses (−25.9 ± 1.6‰), liverworts (−26.3 ± 0.5‰) and algae (−26.3 ± 1.2‰), partly related to habitats controlled by water availability. The δ¹⁵N values of moss samples range widest (−9.0 to 22.8‰, with an average of 4.6 ± 6.6‰). Lichens are on the average most depleted in ¹⁵N (mean = −7.4 ± 6.4‰), whereas algae are most enriched in ¹⁵N (10.0 ± 3.3‰). The broad range of nitrogen isotope compositions suggest that the N source for these Antarctic terrestrial plants is spatially much variable, with the local presence of seabird colonies being particularly significant.     

Diurnal variations of needle water isotopic ratios in two pine species

Diurnal fluctuations of leaf water isotope ratios (δ¹⁸O and δD) were measured for Jeffrey (Pinus jeffreyi Balf.) and lodgepole (Pinus contorta Douglas ex Louden) pine. Two trees per species were sampled every few hours on 15–16 October 2005 and 19–20 June 2006. Diurnal gas exchange was measured during the summer sampling. In fall 2005, leaf water δ¹⁸O ranged from 0.7 to 9.0‰, and leaf water δD ranged from −70 to −50‰. In summer 2006, leaf water δ¹⁸O ranged from 7.7 to 20.7‰, and leaf water δD ranged from −61 to −24‰. Diurnal variation of leaf water isotope values typically reached a maximum in early afternoon, began decreasing around midnight, and reached a minimum in mid-morning. Both periods showed a high degree of enrichment relative to source water, with leaf water–source water enrichments ranging up to 37.8‰ for δ¹⁸O, and up to 95‰ for δD. Leaf water enrichment varied by season with summer enrichment being greater than fall enrichment. A steady-state model (i.e., modified Craig–Gordon modeling) for leaf water isotope compositions did not provide a good fit to measured values of leaf water. In summer, a non-steady state model provided a better fit to the measured data than the steady-state model. Our findings demonstrate substantial leaf water enrichment above source water and diurnal variations in the isotopic composition of leaf water, which has application to understanding short-term variability of atmospheric gases (water vapor, CO₂, O₂), climate studies based on the isotopic composition of tree rings, and ecosystem water fluxes.     

Temporal and spatial partitioning of water resources among eight woody species in a Hawaiian dry forest

Lowland dry forests are unique in Hawaii for their high diversity of tree species compared with wet forests. We characterized spatial and temporal partitioning of soil water resources among seven indigenous and one invasive dry forest species to determine whether the degree of partitioning was consistent with the relatively high species richness in these forests. Patterns of water utilization were inferred from stable hydrogen isotope ratios (δD) of soil and xylem water, zones of soil water depletion, plant water status, leaf phenology, and spatial patterns of species distribution. Soil water δD values ranged from –20‰ near the surface to –48‰ at 130 cm depth. Metrosideros polymorpha, an evergreen species, and Reynoldsia sandwicensis, a drought-deciduous species, had xylem sap δD values of about –52‰, and appeared to obtain their water largely from deeper soil layers. The remaining six species had xylem δD values ranging from –33 to –42‰, and apparently obtained water from shallower soil layers. Xylem water δD values were negatively correlated with minimum annual leaf water potential and positively correlated with leaf solute content, an integrated measure of leaf water deficit. Seasonal patterns of leaf production ranged from dry season deciduous at one extreme to evergreen with near constant leaf expansion rates at the other. Species tapping water more actively from deeper soil layers tended to exhibit larger seasonality of leaf production than species relying on shallower soil water sources. Individuals of Myoporum sandwicense were more spatially isolated than would be expected by chance. Even though this species apparently extracted water primarily from shallow soil layers, as indicated by its xylem δD values, its nearly constant growth rates across all seasons may have been the result of a larger volume of soil water available per individual. The two dominant species, Diospyros sandwicensis and Nestegis sandwicensis, exhibited low leaf water potentials during the dry season and apparently drew water mostly from the upper portion of the soil profile, which may have allowed them to exploit light precipitation events more effectively than the more deeply rooted species. Character displacement in spatial and temporal patterns of soil water uptake was consistent with the relatively high diversity of woody species in Hawaiian dry forests. Received: 20 May 1999 / Accepted: 2 March 2000     

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     

Ontogenetic dietary shifts in European common frog (Rana temporaria) revealed by stable isotopes

During migrations and ontogeny amphibians change their habitat and feeding, and thus are important in linking terrestrial and aquatic ecosystems. We measured δ ¹³C and δ ¹⁵N values of early stages (egg, embryo, tadpole) and toes of adult frogs Rana temporaria, collected from a small wetland in Lithuania. We compared the isotopic composition of these tissues with potential food sources, excrements of tadpoles, and filled intestinal tracts. We found that δ ¹³C values in R. temporaria tadpoles were markedly depleted in comparison to adults, eggs or embryos, demonstrating a terrestrial to aquatic shift in energy sources. After the onset of feeding, tadpoles approached isotopic equilibrium with available food (algae and litter). Tadpoles had higher δ ¹⁵N than both algae and litter, differing by 3.6 and 2.4‰, respectively, and similar δ ¹³C to these sources. However, tadpole excrements and body tissue diverged, with mean δ ¹³C values of excrements (−30.3 ± 1.6‰ SD) more similar to litter (−31.7 ± 1.2‰ SD) and body tissue δ ¹³C (−34.8 ± 0.7‰ SD) more similar to algae (−34.2 ± 4.1‰ SD). This suggests that algal resources are critical in early life stages of this anuran, particularly at stages characterized by high growth and low development (stages: 25–35).     

Challenges using stable isotopes for estimating trophic levels in marine amphipods

In marine food web studies, stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) are widely used to estimate organisms’ trophic levels (TL) and carbon sources, respectively. For smaller organisms, whole specimens are commonly analyzed. However, this “bulk method” involves several pitfalls since different tissues may fractionate stable isotopes differently. We compared the δ¹⁵N and δ¹³C values of exoskeleton versus soft tissue, in relation to whole specimens, of three common Arctic amphipods in Svalbard waters: the benthic Anonyx nugax, the sympagic (ice-associated) Gammarus wilkitzkii and the pelagic Themisto libellula. The δ¹⁵N values of the exoskeletons were significantly lower than those of the soft tissues for A. nugax (10.5 ± 0.7‰ vs. 15.7 ± 0.7‰), G. wilkitzkii (3.3 ± 0.3‰ vs. 8.3 ± 0.4‰) and T. libellula (8.6 ± 0.3‰ vs.10.8 ± 0.3‰). The differences in δ¹³C values between exoskeletons and soft tissues were insignificant, except for A. nugax (−21.2 ± 0.2‰ vs. −20.3 ± 0.2‰, respectively). The δ¹⁵N values of whole organisms were between those of the exoskeletons and the soft tissues, being similarly enriched in ¹⁵N as the exoskeletons (except G. wilkitzkii) and depleted in ¹⁵N by 1.2–3.7‰ compared to the soft tissues. The δ¹⁵N-derived TLs of the soft tissues agreed best with the known feeding preferences of the three amphipods, which suggest a potential underestimation of 0.5–1.0 TL when stable isotope analyses are performed on whole crustaceans with thick exoskeletons. The insignificant or small differences in δ¹³C values among exoskeletons, soft tissues and whole specimens, however, suggest low probability for misinterpretations of crustaceans’ primary carbon source in marine ecosystems with distinctly different δ¹³C-carbon sources.     

δ13C analysis of phloem sap carbon: novel means of evaluating seasonal water stress and interpreting carbon isotope signatures of foliage and trunk wood of Eucalyptus globulus

A recently described phloem-bleeding technique was used to study seasonal changes in δ¹³C, sugar levels and the amino acid:sugar balance of phloem translocate of 2- to 3-year old trees of Eucalyptus globulus at a rain-fed site (Eulup) and a waste-effluent-irrigated site (Albany) in south-west Australia. δ¹³C of phloem sap from the Eulup site fluctuated widely between winter (−27.6‰) and peak summer stress (−20.2‰), compared with a much smaller range of −28.4 to −26.3 at Albany. Seasonal changes in sugar concentrations in sap fluctuated closely with those of phloem δ¹³C, with highest concentrations and least negative δ¹³C values at times of greatest soil water deficit. Molar ratios of amino acids to sugars in phloem sap were similar between plantations in winter through to early summer. They then remained high at the nitrogen-rich effluent-treated site, but fell dramatically once soils dried out at Eulup. Mature leaf dry matter sampled at peak yearly stress (early autumn) showed more negative δ¹³C values than concurrently harvested phloem sap or recently initiated shoot apex dry matter, presumably because the sampled foliage had laid down its structural carbon earlier under relatively unstressed winter/spring conditions. Differences between Albany and Eulup were much greater for δ¹³C of phloem and new apical dry matter than for dry matter of mature foliage. Comparisons of δ¹³C signatures of phloem sap carbon with those of dry matter of nascent xylem tissues showed seasonal fluctuations in δ¹³C of phloem translocate which were mirrored a month or so later by those for xylem carbon. δ¹³C analyses of trunk growth rings from Eulup and Albany showed well-defined seasonal oscillations over the first 2 or 3 years of growth until irrigation commenced at Albany. Fluctuations in δ¹³C at the latter site then became noticeably less pronounced than at Eulup. Future use of phloem sap δ¹³C and solute analyses for studying seasonal water and nutrient status of E. globulus is discussed. Received: 9 April 1998 / Accepted: 20 August 1998     

Functional diversity in an Amazonian rainforest of French Guyana: a dual isotope approach (δ15N and δ13C)

Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2′N, annual precipitation 2200 mm). We assessed leaf δ¹⁵N as a presumptive indicator of symbiotic N₂ fixation, and leaf and wood cellulose δ¹³C as an indicator of leaf intrinsic water-use efficiency (CO₂ assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N₂ fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ¹⁵N was spatially quite uniform in the forest (range 3–5‰), whereas average leaf δ¹⁵N ranged from −0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g⁻¹. Although nodulated trees displayed significantly lower leaf δ¹⁵N values than non-nodulated trees, leaf δ¹⁵N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ¹⁵N values for nodulated and non-nodulated species at the lower end of the δ¹⁵N range. Perturbation did not markedly affect the difference δ¹⁵N_soil−δ¹⁵N_leaf, and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ¹³C were observed in the forest (average values from −31.4 to −26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO₂ assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ¹³C was positively related to total leaf δ¹³C, the former values being 2–3‰ higher than the latter ones. Leaf δ¹³C was not related to leaf δ¹⁵N at either intraspecific or interspecific levels. δ¹³C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ¹³C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species. Received: 18 August 1997 / Accepted: 14 April 1998