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.     

Tracing water uptake by jarrah (Eucalyptus marginata) trees using natural abundances of deuterium

 Seasonal change in the δ²H content of water from twig sap, soil, rainfall and groundwater were measured to determine the water sources accessed by jarrah (Eucalyptus marginata) trees at three sites in Western Australia with differing soils and depths to water table. During winter and spring the main contributor to the water uptake of the trees was stored water in the surface layers of the soil replenished by predominantly winter rainfall. With the onset of summer drought jarrah became more reliant on water from deeper down the profile. There was no clear evidence that jarrah could tap water from groundwater more than 14 m deep in deep sands. Defining the source of water for trees in deep lateritic soils using stable isotopes is hampered by the uniform deuterium profiles down most of the unsaturated zone and into the groundwater. There was a limited response in the δ²H values of sapwater in twigs to changes in the δ²H of the upper layers of the deep sand following input of rainfall in autumn. The damped response was related to the small variation in the δ²H composition of rainfall in most events during the year and the mixing in the tree of water extracted from different locations in the soil profile. Received: 21 August 1995 / Accepted: 3 December 1995     

Hydrogen isotope response to changing salinity and rainfall in Australian mangroves

Hydrogen isotope ratios (²H/¹H, δ²H) of leaf waxes covary with those in precipitation and are therefore a useful paleohydrologic proxy. Mangroves are an exception to this relationship because their δ²H values are also influenced by salinity. The mechanisms underlying this response were investigated by measuring leaf lipid δ²H and leaf and xylem water δ²H and δ¹⁸O values from three mangrove species over 9.5 months in a subtropical Australian estuary. Net ²H/¹H fractionation between surface water and leaf lipids decreased by 0.5–1.0‰ ppt^?1 for n‐alkanes and 0.4–0.8‰ ppt^?1 for isoprenoids. Xylem water was ²H depleted relative to surface water, reflecting ²H discrimination of 4–10‰ during water uptake at all salinities and opportunistic uptake of freshwater at high salinity. However, leaf water ²H enrichment relative to estuary water was insensitive to salinity and identical for all species. Therefore, variations in leaf and xylem water δ²H values cannot explain the salinity‐dependent ²H depletion in leaf lipids, nor the 30‰ range in leaf lipid δ²H values among species. Biochemical changes in direct response to salt stress, such as increased compatible solute production or preferential use of stored carbohydrates, and/or the timing of lipid production and subsequent turnover rates, are more likely causes.     

Effects of leaf water evaporative 2H‐enrichment and biosynthetic fractionation on leaf wax n‐alkane δ2H values in C3 and C4 grasses

Leaf wax n‐alkane δ²H values carry important information about environmental and ecophysiological processes in plants. However, the physiological and biochemical drivers that shape leaf wax n‐alkane δ²H values are not completely understood. It is particularly unclear why n‐alkanes in grasses are typically ²H‐depleted compared with plants from other taxonomic groups such as dicotyledonous plants and why C3 grasses are ²H‐depleted compared with C4 grasses. To resolve these uncertainties, we quantified the effects of leaf water evaporative ²H‐enrichment and biosynthetic hydrogen isotope fractionation on n‐alkane δ²H values for a range of C3 and C4 grasses grown in climate‐controlled chambers. We found that only a fraction of leaf water evaporative ²H‐enrichment is imprinted on the leaf wax n‐alkane δ²H values in grasses. This is interesting, as previous studies have shown in dicotyledonous plants a nearly complete transfer of this ²H‐enrichment to the n‐alkane δ²H values. We thus infer that the typically observed ²H‐depletion of n‐alkanes in grasses (as opposed to dicots) is because only a fraction of the leaf water evaporative ²H‐enrichment is imprinted on the δ²H values. Our experiments also show that differences in n‐alkane δ²H values between C3 and C4 grasses are largely the result of systematic differences in biosynthetic fractionation between these two plant groups, which was on average ?198‰ and?159‰ for C3 and C4 grasses, respectively.     

Changes in wood and foliar δ13C in sugar maple at Gatineau Park,Quebec, Canada

Leaf samples and tree rings formed between the mid‐1960s and mid‐1990s from sugar maple (Acer saccharum Marsh.) at Gatineau Park (45°30′ N, 75°54′ W), Quebec were analysed for δ¹³C. Leaf samples were collected at ground level (1–2 m above ground) at monthly intervals during the summer, whereas tree cores were extracted from the largest trees (d.b.h. > 30 cm) in the young deciduous forest in August 1998. Significant linear decreases in δ¹³C over time were found in foliage and tree rings, but the decrease in δ¹³C was significantly greater in foliage than in the wood. The apparent isotopic discrimination (Δ) of tree rings varied insignificantly around a mean of 18‰, whereas foliar Δ increased significantly from 19‰ in the 1960s to around 23‰ by the mid 1990s, likely as a result of an increasing canopy effect as the forest matured. Using models of carbon discrimination and Δ‐values of the tree rings, we calculate that the intrinsic water use efficiency of mature sugar maple has increased by approximately 4% over the study period.     

Resistance to water flow in xylem of Picea abies (L.) Karst. trees grown under contrasting light conditions

Summary The relative hydraulic conductivity (k) of xylem and resistance (R) to water flow through trunk, primary roots and branches in Picea abies trees growing under contrasting light conditions were investigated. The xylem permeability to water was measured by forcing 10 mM water solution of KC1 through excised wood specimens. From the values of k, the sapwood transverse area and the length of conducting segments, R of the whole trunk, branches and roots was calculated. The relative conductivity of xylem in open-grown trees exceeded that of shade-grown trees by 1.4–3.1 times, while k was closely correlated with the hydraulically effective radius (R _e) of the largest tracheids (R ² was 0.85–0.94 for open- and 0.51–0.79 for shade-grown trees). Because of both a low k and a smaller sapwood area in shade-grown trees the resistance to water movement through their trunk, roots and branches was many times higher. The distribution of R between single segments of the water-conducting pathway differed considerably in trees from different sites. At high water status the largest share of the total resistance in open- as well as shade-grown trees resides in the apical part of the trunk. The contribution of the branches to total xylem resistance is supposed to increase with developing water deficit.     

Spatial,ontogenetic and interspecific variability in stable isotope ratios of nitrogen and carbon of Merluccius capensis and Merluccius paradoxus off South Africa

General linear models (GLMs) were used to determine the relative importance of interspecific, ontogenetic and spatial effects in explaining variability in stable isotope ratios of nitrogen (δ¹⁵N) and carbon (δ¹³C) of the co‐occurring Cape hakes Merluccius capensis and Merluccius paradoxus off South Africa. Significant GLMs were derived for both isotopes, explaining 74 and 56% of observed variance in Merluccius spp. δ¹⁵N and δ¹³C, respectively. Spatial effects (west or south coast) contributed most towards explaining variability in the δ¹⁵N model, with Merluccius spp. off the west coast having higher (by c. 1·4‰) δ¹⁵N levels than Merluccius spp. off the south coast. Fish size and species were also significant in explaining variability in δ¹⁵N, with both species showing significant linear increases in δ¹⁵N with size and M. capensis having higher (by c. 0·7‰) δ¹⁵N values than M. paradoxus. Species and coast explained most and similar amounts of variability in the δ¹³C model, with M. capensis having higher (by c. 0·8‰) δ¹³C values than M. paradoxus, and values being lower (by c. 0·7‰) for fishes off the west coast compared with the south coast. These results not only corroborate the knowledge of Merluccius spp. feeding ecology gained from dietary studies, in particular the ontogenetic change in trophic level corresponding to a changing diet, but also that M. capensis feeds at a slightly higher trophic level than M. paradoxus. The spatial difference in Merluccius spp. δ¹⁵N appears due to a difference in isotopic baseline, and not as a result of Merluccius spp. feeding higher in the food web off the west than the south coast, and provides new evidence that corroborates previous observations of biogeographic differences in isotopic baselines around the South African coast. This study also provides quantitative data on the relative trophic level and trophic width of Cape hakes over a large size range that can be used in ecosystem models of the southern Benguela.     

Feeding ecology of pelagic fish larvae and juveniles in slope waters of the Gulf of Mexico

Stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) were used to investigate feeding patterns of larval and early juvenile pelagic fishes in slope waters of the Gulf of Mexico. Contribution of organic matter supplied to fishes and trophic position within this pelagic food web was estimated in 2007 and 2008 by comparing dietary signatures of the two main producers in this ecosystem: phytoplankton [based on particulate organic matter (POM)] and Sargassum spp. Stable isotope ratios of POM and pelagic Sargassum spp. were significantly different from one another with δ¹³C values of POM depleted by 3–6‰ and δ¹⁵N values enriched by 2 relative to Sargassum spp. Stable isotope ratios were significantly different among the five pelagic fishes examined: blue marlin Makaira nigricans, dolphinfish Coryphaena hippurus, pompano dolphinfish Coryphaena equiselis, sailfish Istiophorus platypterus and swordfish Xiphias gladius. Mean δ¹³C values ranged almost 2 among fishes and were most depleted in I. platypterus. In addition, mean δ¹⁵N values ranged 4–5 with highest mean values found for both C. hippurus and C. equiselis and the lowest mean value for M. nigricans during both years. Increasing δ¹³C or δ¹⁵N with standard length suggested that shifts in trophic position and diet occurred during early life for several species examined. Results of a two‐source mixing model suggest approximately an equal contribution of organic matter by both sources (POM = 55%; pelagic Sargassum spp. = 45%) to the early life stages of pelagic fishes examined. Contribution of organic matter, however, varied among species, and sensitivity analyses indicated that organic source estimates changed from 2 to 13% for a δ¹³C fractionation change of ±0·25‰ or a δ¹⁵N fractionation change of ± 1·0‰ relative to original fractionation values.     

Aerial decay influence on the stable oxygen and carbon isotope ratios in tree ring cellulose

Sub-fossil wood is often affected by the decaying process that introduces uncertainties in the measurement of oxygen and carbon stable isotope composition in cellulose. Although the cellulose stable isotopes are widely used as climatic proxies, our understanding of processes controlling their behavior is very limited. We present here a comparative study of stable oxygen and carbon isotope ratios in tree ring cellulose in decayed and non-decayed wood samples of Swiss stone pine (Pinus cembra) trees. The intra-ring stable isotope variability (around the circumference of a single ring) was between 0.1 and 0.5‰ for δ¹⁸O values and between 0.5 and 1.6‰ for δ¹³C values for both decayed and non-decayed wood. Observed intra-tree δ¹⁸O variability is less than that reported in the literature (0.5–1.5‰), however, for δ¹³C it is larger than the reported values (0.7–1.2‰). The inter-tree variability for non-decayed wood ranges between 1.1 and 2.3‰ for δ¹⁸O values, and between 2 and 4.7‰ for δ¹³C values. The inter-tree differences for δ¹⁸O values are similar to those reported in the literature (1–2‰ for oxygen and 1–3‰ for carbon) but are larger for δ¹³C values. We have found that the differences for δ¹⁸O and δ¹³C values between decayed and non-decayed wood are smaller than the variation among different trees from the same site, suggesting that the decayed wood can be used for isotopic paleoclimate research.     

Bulk hydrogen stable isotope composition of seaweeds: Clear separation between Ulvophyceae and other classes

Little is known about the bulk hydrogen stable isotope composition (δ²H) of seaweeds. This study investigated the bulk δ²H in several different seaweed species collected from three different beaches in Brazil, Australia, and Argentina. Here, we show that Ulvophyceae (a group of green algae) had lower δ²H values (between ?94‰ and ?130‰) than red algae (Florideophyceae), brown algae (Phaeophyceae), and species from the class Bryopsidophyceae (another group of green algae). Overall the latter three groups of seaweeds had δ²H values between ?50‰ and ?90‰. These findings were similar at the three different geographic locations. Observed differences in δ²H values were probably related to differences in hydrogen (H) metabolism among algal groups, also observed in the δ²H values of their lipids. The marked difference between the δ²H values of Ulvophyecae and those of the other groups could be useful to trace the food source of food webs in coastal rocky shores, to assess the impacts of green tides on coastal ecosystems, and to help clarify aspects of their phylogeny. However, reference materials for seaweed δ²H are required before the full potential of using the δ²H of seaweeds for ecological studies can be exploited.     

Tracing dietary origins of aphids and the predatory beetle Propylea japonica in agricultural systems using stable isotope analyses

Tracing dietary origins of the predatory beetle Propylea japonica (Thunberg) (Coleoptera: Coccinellidae) aids understanding their roles in the food web and provides information to develop strategies for effective conservation in agroecosystems comprised of wheat [Triticum aestivum L. (Poaceae)], cotton [Hirsutum spp. (Malvaceae)], and maize [Zea mays L. (Poaceae)]. Intrinsic markers of carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) in P. japonica need to be developed to ascertain the source(s) of diet. Experiments were carried out to examine the changes of δ¹³C and δ¹⁵N among the three crops, pests (wheat, cotton, and maize aphids; all Hemiptera: Aphididae), and P. japonica fed on aphids of each of the three crops. Results indicated that δ¹³C values in P. japonica fed on wheat, cotton, and maize aphids were ?27.2 to ?26.5‰, ?24.2 to ?23.9‰, and ?11.0 to ?10.7‰, respectively, whereas their δ¹⁵N values were 1.1 to 2.9‰, 6.0 to 7.4‰, and ?0.6 to 0.1‰, respectively. δ¹³C and δ¹⁵N plots clearly identify the three crops, the dietary origins of the aphids, and the host origins of the aphid prey consumed by the ladybird beetles, as each pathway displays a non‐overlapping pattern. Based on the values of δ¹³C and δ¹⁵N of the three food webs, dietary origins can be traced in the predatory beetle P. japonica derived from wheat, cotton, and maize crops.     

Tracing sources of carbon and hydrogen to stored lipids in the migratory moth,Mythimna unipuncta using stable isotopes (δ2H, δ13C)

Stable isotope measurements of insect tissues can be used to determine origin and migratory patterns. The isotopic links between diet and stored lipids in laboratory reared true armyworm moths (Mythimna unipuncta, Haw.) were investigated using δ¹³C and δ²H measurements. Newly emerged moths were fed synthetic nectars, consisting of different carbohydrate and water sources that were isotopically distinct from those in the larval diet. After 4 days of feeding, insects were sacrificed, and fat‐body lipids were extracted for isotopic analysis. When held on a constant nectar source, adult diet contributed ～87% of the C and 44% H of lipids. For H, 27% and ～ 17% of lipid was derived from carbohydrates and water, respectively, with the remainder presumably from the larval stage. When the isotopic composition of the nectar source was switched there was rapid and exponential temporal change in lipid isotopic profiles. The relevance of our findings to tracing origins of stored lipids in migrant Lepidoptera is discussed.     

Carbon (δ13C) and nitrogen (δ15N) stable isotope composition in plant and soil in Southern Patagonia's native forests

Stable isotope natural abundance measurements integrate across several biogeochemical processes in ecosystem N and C dynamics. Here, we report trends in natural isotope abundance (δ¹³C and δ¹⁵N in plant and soil) along a climosequence of 33 Nothofagus forest stands located within Patagonia, Southern Argentina. We measured 28 different abiotic variables (both climatic variables and soil properties) to characterize environmental conditions at each of the 33 sites. Foliar δ¹³C values ranged from ?35.4‰ to ?27.7‰, and correlated positively with foliar δ¹⁵N values, ranging from ?3.7‰ to 5.2‰. Soil δ¹³C and δ¹⁵N values reflected the isotopic trends of the foliar tissues and ranged from ?29.8‰ to ?25.3‰, and ?4.8‰ to 6.4‰, respectively, with no significant differences between Nothofagus species (Nothofagus pumilio, Nothofagus antarctica, Nothofagus betuloides). Principal component analysis and multiple regressions suggested that mainly water availability variables (mean annual precipitation), but not soil properties, explained between 42% and 79% of the variations in foliar and soil δ¹³C and δ¹⁵N natural abundance, which declined with increased moisture supply. We conclude that a decline in water use efficiency at wetter sites promotes both the depletion of heavy C and N isotopes in soil and plant biomass. Soil δ¹³C values were higher than those of the plant tissues and this difference increased as annual precipitation increased. No such differences were apparent when δ¹⁵N values in soil and plant were compared, which indicates that climatic differences contributed more to the overall C balance than to the overall N balance in these forest ecosystems.     

Stipa tenacissima Does not Affect the Foliar δ13C and δ15N of Introduced Shrub Seedlings in a Mediterranean Semi-arid Steppe

Recent studies have shown that the tussock grass Stipa tenacissima L. facilitates the establishment of late-successional shrubs, in what constitutes the first documented case of facilitation of woody plants by grasses. With the aim of increasing our knowledge of this interaction, in the present study we investigated the effects of S. tenacissima on the foliar δ13C, δ15N, nitrogen concentration, and carbon ： nitrogen ratio of introduced seedlings of Pistacia lentiscus L., Quercus coccifera L., and Medicago arborea L. in a semi-arid Mediterranean steppe. Six months after planting, the values of δ13C ranged between -26.9‰ and -29.6‰, whereas those of δ15N ranged between -1.9‰ and 2.7‰. The foliar C ： N ratio ranged between 10.7 and 53.5, and the nitrogen concentration ranged between 1.0% and 4.4%. We found no significant effect of the microsite provided by S. tenacissima on these variables in any of the species evaluated. The values of δ13C were negatively correlated with predawn water potentials in M. arborea and were positively correlated with relative growth rate in Q. coccifera. The values of δ15N were positively correlated with the biomass allocation to roots in the latter species. The present results suggest that the modification of environmental conditions in the are surrounding S. tenacissima was not strong enough to modify the foliar isotopic and nitrogen concentration of shrubs during the early stages after planting.     

The flow of water into fruit trees. I. Resistances to water flow through roots and stems

Relative conductivity (K) to water in healthy apple trees ranged from maximum values of 18.2 cm³.100 s^-1.cm length.0.001 Pas.kPa^-1.cm^-2 xylem area, for major suberized roots to values of 1.6 for 1-yr-old twigs. The values for equivalent parts of healthy cherry trees were 26.3 and 3.3. Trees with roots affected by the larvae of the fruit tree root weevil (Leptopius squalidus) which causes either chronic growth decline or sudden wilting and death, had values as low as 1% of healthy trees, in those parts of the tree showing wilting and lack of growth. Water flow under pressure into the root systems of healthy apple trees increased linearly with increases in pressure from 200 to 800 kPa. Flows into dormant and active root systems respectively were 0.6 and 1.7 cm³.100 s^-1. 100 cm² root surface area. 100 kPa^-1.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Waterlogging and fire impacts on nitrogen availability and utilization in a subtropical wet heathland (wallum)

Protein, amino acids and ammonium were the main forms of soluble soil nitrogen in the soil solution of a subtropical heathland (wallum). After fire, soil ammonium and nitrate increased 90- and 60-fold, respectively. Despite this increase in nitrate availability after fire, wallum species exhibited uniformly low nitrate reductase activities and low leaf and xylem nitrate. During waterlogging soil amino acids increased, particularly γ-aminobutyric acid (GABA) which accounted for over 50% of amino nitrogen. Non-mycorrhizal wallum species were significantly (P < 0.05) ¹⁵N-enriched (0.3–4.3‰) compared to species with mycorrhizal associations (ericoid-type, ecto-, va-mycorrhizal) which were strongly depleted in ¹⁵N (-6.3 to -1.8‰). Lignotubers and roots had δ¹⁵N signatures similar to that of the leaves of respective species. The exceptions were fine roots of ecto-, ecto/va-, and ericoid type mycorrhizal species which were enriched in ¹⁵N (0.1–2.4‰). The 5¹⁵N signatures of δ¹⁵N_{total soil N} and δ¹⁵N_{soil NH4+} were in the range 3.7–4.5‰, whereas δ¹⁵N_{soil NO3?} was significantly (P < 0.05) more enriched in ¹⁵N (9.2–9.8‰). It is proposed that there is discrimination against ¹⁵N during transfer of nitrogen from fungal to plant partner. Roots of selected species incorporated nitrogen sources in the order of preference: ammonium > glycine > nitrate. The exception were proteoid roots of Hakea (Proteaceae) which incorporated equal amounts of glycine and ammonium.     

The effect of fertilization on sap flux and canopy conductance in a Eucalyptus saligna experimental forest

Land devoted to plantation forestry (50 million ha) has been increasing worldwide and the genus Eucalyptus is a popular plantation species (14 million ha) for its rapid growth and ability to grow well on a wide range of sites. Fertilization is a common silvicultural tool to improve tree growth with potential effects on stand water use, but the relationship between wood growth and water use in response to fertilization remains poorly quantified. Our objectives in this study were to determine the extent, timing and longevity of fertilization effects on water use and wood growth in a non‐water limited Eucalyptus saligna experimental forest near Hilo, HI. We evaluated the short‐ and long‐term effects of fertilization on water use by measuring sap flux per unit sapwood area, canopy conductance, transpiration per unit leaf area and water‐use efficiency in control and fertilized stands. Short‐term effects were assessed by comparing sap flux before and after fertilizer application. Long‐term effects were assessed by comparing control plots and plots that had received nutrient additions for 5 years. For the short‐term response, total water use in fertilized plots increased from 265 to 487 mm yr^?1 during the 5 months following fertilization. The increase was driven by an increase in stand leaf area accompanied by an increase in sap flux per unit sapwood area. Sap flux per unit leaf area and canopy conductance did not differ during the 5 months following fertilizer additions. For the last 2 months of our short‐term measurements, fertilized trees used less water per unit carbon gain (361 compared with 751 kg H₂O kg C^?1 in control stands). Trees with 5 years of fertilization also used significantly more water than controls (401 vs. 302 mm yr^?1) because of greater leaf area in the fertilized stands. Sap flux per unit sapwood area, sap flux per unit leaf area, and canopy conductance did not differ between control and fertilized trees in the long‐term plots. In contrast to the short‐term response, the long‐term response of water use per unit wood growth was not significant. Overall, fertilization of E. saligna at our site increased stand water use by increasing leaf area. Fertilized trees grew more wood and used more water, but fertilization did not change wood growth per unit water use.     

Identifying groundwater‐fed climate refugia in remote arid regions with citizen science and isotope hydrology

1. Groundwater and surface water‐fed systems act as biodiversity hotspots and ecological refuges and evolutionary refugia in arid regions. Groundwater‐dominated systems are sustained by underground aquifers that are recharged by rain that has fallen in the distant past, while surface water‐dominated systems are fed by recent local rain or floods. Some waterbodies are fed by a mixture of these sources. Perennial, groundwater‐dominated systems will act as refuges and refugia under future rainfall declines associated with global warming. We sought to identify climate refugia, based on groundwater dominance, by using isotope hydrology to characterise water samples collected by citizen scientists across arid central Australia.
2. There is a linear relationship between hydrogen isotopes (²H/¹H, δ²H) and oxygen isotopes (¹⁸O/¹⁶O, δ¹⁸O) in rainfall. This relationship is known as the meteoric water line (MWL). By comparing our samples with the Australian MWL, and developing a local evaporation line, we were able to test the hypotheses that groundwater‐dominated systems will follow the Australian MWL while temporary systems follow the local evaporation line, and, accordingly, distinguish between groundwater and surface water‐dominated systems. The isotopic composition of samples collected over a 36‐month period was determined using isotope ratio infrared spectrometry. The electrical conductivity of each sample was recorded to determine where freshwater is available for biota within this arid region.
3. Over 240 water samples were collected from 62 waterbodies and seven bores (groundwater wells) spanning an area of more than 250,000 km². Approximately 75% of the samples were collected by citizen scientists and 25% by research scientists.
4. Twenty groundwater‐dominated waterbodies, characterised by a small range of δ²H and δ¹⁸O values (c. ?55 to ?20‰ and c. ?9 to ?3‰, respectively) clustered around the long‐term mean composition of rainfall (δ²H = ?37.5‰, δ¹⁸O = ?6.4‰), were identified as future evolutionary refugia. These sites are likely to contain water through the most severe of droughts and will be critically important for the persistence of water‐dependent species.
5. Based on their isotopic composition, we identified 45 waterbodies (rockholes/waterholes) as temporary or ephemeral (δ²H c. ?40 to ?100‰ and δ¹⁸O c. ?4 to +25‰), that is, with no evidence of groundwater inflow. These, together with waterbodies supported by a mix of groundwater and surface water, can act as stepping stones and form part of the aquatic mosaic that is critical to supporting species in arid regions. Over two‐thirds of the waterholes sampled were very fresh (electrical conductivity <0.8 mS/cm), indicating that they provide the freshwater needed to support much of the regional aquatic and terrestrial fauna.
6. All evolutionary refugia are located within protected areas (i.e. national parks or Indigenous Protected Areas), but some are subject to the impacts of feral animal species and invasive plants. Our findings indicate where control programmes and restoration actions can be prioritised to support biodiversity conservation and climate change adaptation. Our approach, combining citizen science and isotope hydrology, can be used to identify future refugia in other remote and arid regions where water scarcity is likely to increase under global climate change.

     

Using Soil 13C to Detect the Historic Presence of Schizachyrium scoparium (Little Bluestem) Grasslands on Martha's Vineyard

Abstract We used differences in soil carbon δ¹³C values between forested sites and grasslands dominated by the C₄ grass Schizachyrium scoparium (little bluestem) to detect the presence of former grasslands in the historical landscape of the coastal sand plain of Martha's Vineyard, Massachusetts, U.S.A. Soil δ¹³C was measured at (1) sites with long‐term forest or grassland vegetation and (2) sites with known histories where forest vegetation invaded grassland and where forest converted to grassland. The δ¹³C of soil under long‐term grassland was –24.1‰ at 0 to 2 cm depth and –23.4‰ at 2 to 10 cm and was enriched by 3.4‰ and 2.8‰ compared with soil under long‐term forest. In forests that invaded grasslands dominated by S. scoparium, soil δ¹³C decreased as C derived from trees replaced C from S. scoparium. This decline occurred faster in surface soils and in the light soil organic matter fraction than in the mineral soil. In forests that converted to grasslands, soil δ¹³C increased and the rate of increase was similar in surface and mineral soil and in the different soil organic matter fractions. Rates of change indicated that soil δ¹³C could be used to detect changes in vegetation involving the presence or absence of S. scoparium during the last 150 years. Application of this model to a potential grassland restoration site on Martha's Vineyard where the landscape history was not known indicated that the site was previously unoccupied by S. scoparium during this time. The δ¹³C of surface mineral soil can be useful for detecting the presence of historic S. scoparium grasslands but only in the period well after European settlement of these coastal sand plain landscapes.