Changes in Canopy Processes Following Whole-Forest Canopy Nitrogen Fertilization of a Mature Spruce-Hemlock Forest

Abstract Most experimental additions of nitrogen to forest ecosystems apply the N to the forest floor, bypassing important processes taking place in the canopy, including canopy retention of N and/or conversion of N from one form to another. To quantify these processes, we carried out a large-scale experiment and determined the fate of nitrogen applied directly to a mature coniferous forest canopy in central Maine (18–20 kg N ha⁻¹ y⁻¹ as NH₄NO₃ applied as a mist using a helicopter). In 2003 and 2004 we measured NO₃ ⁻, NH₄ ⁺, and total dissolved N (TDN) in canopy throughfall (TF) and stemflow (SF) events after each of two growing season applications. Dissolved organic N (DON) was greater than 80% of the TDN under ambient inputs; however NO₃ ⁻ accounted for more than 50% of TF N in the treated plots, followed by NH₄ ⁺ (35%) and DON (15%). Although NO₃ ⁻ was slightly more efficiently retained by the canopy under ambient inputs, canopy retention of NH₄ ⁺as a percent of inputs increased markedly under fertilization. Recovery of less than 30% of the fertilizer N in TF suggested that the forest canopy retained more than 70% of the applied N (>80% when corrected for N which bypassed tree surfaces at the time of fertilizer addition). Results from plots receiving ¹⁵N enriched NO₃ ⁻ and NH₄ ⁺ confirmed bulk N estimations that more NO₃ ⁻ than NH₄ ⁺ was washed from the canopy by wet deposition. The isotope data did not show evidence of canopy nitrification, as has been reported in other spruce forests receiving much higher N inputs. Conversions of fertilizer-N to DON were observed in TF for both ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ additions, and occurred within days of the application. Subsequent rain events were not significantly enriched in ¹⁵N, suggesting that canopy DON formation was a rapid process related to recent N inputs to the canopy. We speculate that DON may arise from lichen and/or microbial N cycling rather than assimilation and re-release by tree tissues in this forest. Canopy retention of experimentally added N may meet and exceed calculated annual forest tree demand, although we do not know what fraction of retained N was actually physiologically assimilated by the plants. The observed retention and transformation of DIN within the canopy demonstrate that the fate and ecosystem consequences of N inputs from atmospheric deposition are likely influenced by forest canopy processes, which should be considered in N addition studies. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Large Canopy Exchange Fluxes of Inorganic and Organic Nitrogen and Preferential Retention of Nitrogen by Epiphytes in a Tropical Lowland Rainforest

Little is known about how tropical forest canopies interact with atmospheric nitrogen deposition and how this affects the internal nutrient dynamics and the processing of external nutrient inputs. The objectives of this study therefore were (1) to investigate gross and net canopy nitrogen (N) fluxes (retention and leaching) and (2) the effect of canopy components on net canopy N retention. Tracers were applied on detached branches in a tropical wet lowland rainforest, Costa Rica. A novel ¹⁵N pool dilution method showed that gross canopy fluxes (retention and leaching) of NO₃ ^?, NH₄ ⁺, and dissolved organic nitrogen (DON) were remarkably higher than net throughfall fluxes. Gross fluxes of NH₄ ⁺ and NO₃ ^? resulted in a negligible net flux whereas DON showed net uptake by the canopy. The highest quantity of ¹⁵N was recovered in epiphytic bryophytes (16.4%) although the largest biomass fraction was made up of leaves. The study demonstrates that tracer applications allow investigation of the dynamic and complex canopy exchange processes and that epiphytic communities play a major role in solute fluxes in tree canopies and therefore in the nutrient dynamics of tropical rain forests.     

Spatial interpolation of N concentrations and δ15N values in the moss Hypnum cupressiforme collected in the forests of Slovenia

Mosses were collected from 103 locations and analyzed for N concentration and δ¹⁵N values. At each location, the samples were collected from two types of site: under the tree canopies and in the adjacent forest openings. A spatial correlation was detected, and ordinary kriging could be applied only to the N in the mosses that were collected in the openings. In contrast, a spatial correlation was not detected for N in the mosses that were collected under the canopies or for the δ¹⁵N of the mosses at either type of site. For those, the spatial interpolation was calculated as the sum of the regression predictions and the inverse distance weighted interpolation of residuals. The maps for the N at both site types (canopy/open) identified similar areas with increased N concentrations. The sole exception was the mosses collected under the canopy, for which some local emitters of NO_x were also identified.     

Influences of elemental interactions and pedogenic processes in organic matter dynamics

This paper reviews progress in understanding the processes which are important in elemental interactions and which influence organic matter composition of soils of the Great Plains in N. America. Comparison of grassland (semiarid) soils along environmental gradients and cultivation chrono- and toposequences with adjacent forest (subhumid) soils and consideration of the C/N/P/S ratios of organic matter of genetic horizons in the solum have emphasized the importance of movement of low molecular weight organic compounds in soil solution in addition to microbial degradation in the formulation of organic matter in soils. Phosphorus forms and transformations help to provide both an index on weathering and insight into textural influences. Use of ¹⁵N and ³⁴S in combination with¹⁴C and other radioisotopes has provided valuable information on processes. Submicroscopy techniques in combination with cytoplasmic staining techniques have focussed attention in a realistic way on the mechanisms of organic matter stability. More attention must be paid to the catalytic role of soil inorganic constituents and selected minerals in the abiotic formation of stable organic matter. Conceptual and mathematical simulation models have an invaluable role in focussing attention on important processes and verifying hypotheses.     

A break in the nitrogen cycle in aridlands? Evidence from δp15N of soils

We examined the content and isotopic composition of nitrogen within soils of a juniper woodland and found that a cryptobiotic crust composed of cyanobacteria, lichens, and mosses was the predominant source of nitrogen for this ecosystem. Disturbance of the crust has resulted in considerable spatial variability in soil nitrogen content and isotopic composition; intercanopy soils were significantly depleted in nitrogen and had greater abundance of ¹⁵N compared to intra-canopy soils. Variations in the ¹⁵N/¹⁴N ratio for inter- and intra-canopy locations followed similar Rayleigh distillation curves, indicating that the greater ¹⁵N/¹⁴N ratios for inter-canopy soils were due to relatively greater net nitrogen loss. Coverage of cryptobiotic crusts has been reduced by anthropogenic activities during the past century, and our results suggest that destruction of the cryptobiotic crust may ultimately result in ecosystem degradation through elimination of the predominant source of nitrogen input.     

Does canopy nitrogen uptake enhance carbon sequestration by trees?

Temperate forest ¹⁵N isotope trace experiments find nitrogen (N) addition‐driven carbon (C) uptake is modest as little additional N is acquired by trees; however, several correlations of ambient N deposition against forest productivity imply a greater effect of atmospheric nitrogen deposition than these studies. We asked whether N deposition experiments adequately represent all processes found in ambient conditions. In particular, experiments typically apply ¹⁵N to directly to forest floors, assuming uptake of nitrogen intercepted by canopies (CNU) is minimal. Additionally, conventional ¹⁵N additions typically trace mineral ¹⁵N additions rather than litter N recycling and may increase total N inputs above ambient levels. To test the importance of CNU and recycled N to tree nutrition, we conducted a mesocosm experiment, applying 54 g N/¹⁵N ha^?1 yr^?1 to Sitka spruce saplings. We compared tree and soil ¹⁵N recovery among treatments where enrichment was due to either (1) a ¹⁵N‐enriched litter layer, or mineral ¹⁵N additions to (2) the soil or (3) the canopy. We found that 60% of ¹⁵N applied to the canopy was recovered above ground (in needles, stem and branches) while only 21% of ¹⁵N applied to the soil was found in these pools. ¹⁵N recovery from litter was low and highly variable. ¹⁵N partitioning among biomass pools and age classes also differed among treatments, with twice as much ¹⁵N found in woody biomass when deposited on the canopy than soil. Stoichiometrically calculated N effect on C uptake from ¹⁵N applied to the soil, scaled to real‐world conditions, was 43 kg C kg N^?1, similar to manipulation studies. The effect from the canopy treatment was 114 kg C kg N^?1. Canopy treatments may be critical to accurately represent N deposition in the field and may address the discrepancy between manipulative and correlative studies.     

Isotopic evidence for the occurrence of biological nitrification and nitrogen deposition processing in forest canopies

This study examines the role of tree canopies in processing atmospheric nitrogen (N_dep) for four forests in the United Kingdom subjected to different N_dep: Scots pine and beech stands under high N_dep (HN, 13–19 kg N ha^?1 yr^?1), compared to Scots pine and beech stands under low N_dep (LN, 9 kg N ha^?1 yr^?1). Changes of NO₃‐N and NH₄‐N concentrations in rainfall (RF) and throughfall (TF) together with a quadruple isotope approach, which combines δ¹⁸O, Δ¹⁷O and δ¹⁵N in NO₃^? and δ¹⁵N in NH₄⁺, were used to assess N transformations by the canopies. Generally, HN sites showed higher NH₄‐N and NO₃‐N concentrations in RF compared to the LN sites. Similar values of δ¹⁵N‐NO₃^? and δ¹⁸O in RF suggested similar source of atmospheric NO₃^? (i.e. local traffic), while more positive values for δ¹⁵N‐NH₄⁺ at HN compared to LN likely reflected the contribution of dry NH_x deposition from intensive local farming. The isotopic signatures of the N‐forms changed after interacting with tree canopies. Indeed, ¹⁵N‐enriched NH₄⁺ in TF compared to RF at all sites suggested that canopies played an important role in buffering dry N_dep also at the low N_dep site. Using two independent methods, based on δ¹⁸O and Δ¹⁷O, we quantified for the first time the proportion of NO₃^? in TF, which derived from nitrification occurring in tree canopies at the HN site. Specifically, for Scots pine, all the considered isotope approaches detected biological nitrification. By contrast for the beech, only using the mixing model with Δ¹⁷O, we were able to depict the occurrence of nitrification within canopies. Our study suggests that tree canopies play an active role in the N cycling within forest ecosystems. Processing of N_dep within canopies should not be neglected and needs further exploration, with the combination of multiple isotope tracers, with particular reference to Δ¹⁷O.     

Stable isotopes as indicators of wastewater effects on the macroinvertebrates of urban rivers

Rivers in urban locations frequently receive contaminated wastewater and particulate waste either directly from storm overflows or from sewage treatment facilities. Although many urban streams are now recovering from wide-scale historic pollution, lower-level effects on water chemistry, nutrients and biotic composition are still widespread. We aimed to determine whether such effects could be detected using stable isotope ratios (??¹⁵N, ??¹³C and ??³⁴S) in macroinvertebrates alone or in conjunction with traditional biomonitoring. Macroinvertebrates were collected upstream and downstream of 11 different secondary wastewater treatment works (WwTW) in South Wales and the Welsh borders (United Kingdom). Overall, mean invertebrate ??¹⁵N signatures downstream of the WwTW were significantly enriched despite variation amongst sites. Moreover, changes between upstream and downstream macroinvertebrate ??¹⁵N values were highly correlated with patterns in macroinvertebrate community composition, increased total macroinvertebrate abundance, and reduced Shannon Diversity and other biomonitoring indices (% EPT, % shredders and ASPT scores). Changes in invertebrate ??¹⁵N values also paralleled the consented discharge volumes and population equivalents from each WwTW. In contrast, isotopic ratios of ??¹³C and ??³⁴S were unable to distinguish or quantify wastewater input into the rivers but differences were apparent amongst study streams. Overall, these results suggest that macroinvertebrate ??¹⁵N signatures can detect and quantify the effects of secondary sewage treatment inputs to riverine ecosystems. Moreover, the method potentially provides a sensitive means for tracing sewage-derived nutrients into food webs while inferring effects on aquatic communities where sewage-loads are subtle or confounded by other stressors.     

Sulfur,carbon, and nitrogen stable isotope ratios in soft tissues and trophic relationships of fish from the near-shore waters of the peter the great bay in the Sea of Japan

Stable isotope ratios of sulfur (³⁴S/³²S), carbon (¹³C/¹²C), and nitrogen (¹⁵N/¹⁴N) were analyzed in the soft tissues of 12 common species of fish from the near-shore waters of the Peter the Great Bay in the Sea of Japan. The average δ¹³C values of individual species varied from −20.7‰ for planktivorous fish to −16.8‰ for benthivorous fish, reflecting the growing relative contribution of benthic primary producers to fish nutrition. The majority of the various species representatives studied can be assigned to one trophic level, as indicated by their narrow range of δ¹⁵N values (9.9 to 12.6‰). Large interspecific variations were found in the sulfur stable isotope ratios of fish (the mean δ³⁴S values ranged from 11.2 to 19.5‰). This is the result of the different contributions to fish nutrition of infaunal invertebrates that are depleted in ³⁴S due to the microbial food chain of the bottom sediments.     

Exchange of Proton and Major Elements in Two-Layer Canopies Under Acid Rain in a Subtropical Evergreen Forest in Central-South China

Canopy exchanges of H^＋ and N （NH4^＋-N, NO3^--N） and other major ions were evaluated and quantified In twolayer canopies based on throughfall measurements in Shaoshan Forest during the period 2000-2002, central-south China, The collected annual rainfall, throughfall, and sub-throughfall were 1 401, 1 191, and 1 084 mm/year, respectively. Fifteen percent and 8% of rainfall （or 9% of throughfall） were intercepted by the top canopy and sub-canopy layers, respectively, The foliar leaching of base cations from the top canopy was significantly higher than that from the sub-canopy, and the latter accounted for 25% of the former. The uptake of H^＋ and NH4^＋ was significantly higher in the top canopy than in the sub-canopy, indicating that the canopy buffering capacity in the top canopy was stronger than the sub-canopy; Mg^2＋ can be absorbed from water flux on the sub-canopy foliar surfaces to compensate for the Mg deficit in the forest soil during the growing season,     

Intracanopy variation in nitrogen cycling through leaves is influenced by irradiance and proximity to developing fruit in mature walnut trees

Intracanopy variation in net leaf nitrogen (N) resorption and N cycling through leaves in mature walnut (Juglans regia L. cv Hartley) trees were monitored in 3 different years. Differential irradiance among the spurs sampled was inferred from differences among leaves in dry weight per unit area (LW/LA) which varied from 4.0 mg · cm^–2 to 7.0 mg · cm^–2 in shaded (S) and exposed (E) canopy positions, respectively. Our results, using ¹⁵N-depleted (NH₄)₂SO₄ validated the concept that N influx and efflux through fully expanded leaves occurred concurrently during the period of embryo growth. Additionally, it also suggested that N influx into leaves was substantially greater in exposed as compared with shaded canopy positions. Because of its well documented phloem immobility, leaf Ca accumulation was used to better estimate the relative influx of N into exposed and shaded leaves. N cycling varied locally within the tree canopy, i. e. Ca (and presumably N) influx was 100% greater in exposed than shaded tree canopy positions, but influx was not influenced significantly by the proximity of developing fruit. In contrast, both the amount and percentage N efflux was significantly greater during embryo growth in fruit-bearing than defruited spurs. Net leaf N resorption averaged 2–4 times greater (25–30%) in fruit-bearing spurs than the 5–10% decrease in the leaf N content in defruited spurs. Since about 90% of leaf N content reportedly occurs as protein, fruit N demand apparently influenced protein turnover and catalysis in associated spur leaves. The amount of leaf N resorption was greater in exposed than shaded positions in the tree canopy in 2 of the 3 years of data collection. Our data show that like leaf N content, N influx, N efflux and net leaf N resorption vary throughout mature walnut tree canopies under the combined local influences of fruiting and irradiance.     

Understanding the fate and linkage of N and S in earthworm-engineered peat soil by coupling stable isotopes and nano-scale secondary ion mass spectrometry

Earthworm burrow-lining is a biophysical interface where the relationship between the spatial location of active soil microorganisms and the actual physical structure of the soil has a major influence on geochemical processes and nutrient cycling. Isotopic tracers and NanoSIMS were combined to highlight, at the nanometric scale, N and S transfers from earthworms to this non-destructured biophysical interface. Peat mesocosms were inoculated with double labelled earthworms (¹⁵N and ³⁴S; Lumbricus rubellus H.) and then sampled for NanoSIMS quantitative measurements. The maximum enrichments of ¹⁵N and ³⁴S detected in peat after 24 days were 1.6 and 5.6 APE (Atom % excess), respectively. NanoSIMS analyses, performed along a transect in peat at increasing distances from contact between the epidermis of an earthworm and its burrow-lining, revealed rapid and brief transfers of biogenic N and S into peat with two contrasted distributions i.e., decreasing ¹⁵N and patchy ³⁴S. The sphere of influence of L. rubellus was clearly delimited after 24 days at 1 mm around the burrow, thus specifying the functional traits of this epi-anecic earthworm. The observed N inputs into peat were probably derived from oxidation of the labile products of earthworm metabolism, such as urea and/or ammonium, as shown by the strong linkage of ¹⁵N with oxygen measured as ¹⁶O^?. The transfers of labile S products from earthworm to the surrounding soil are reported here for the first time and could be derived from mucoprotein secretion. The interesting spatially inverse relationship between ¹⁵N and ³⁴S enriched-spots at increasing distances from the burrow-lining highlighted different fates of the biogenic N and S compounds excreted by earthworms and could reveal the activities of syntrophic partnerships in N and S cycling. The strategy of combining isotopic tracers with NanoSIMS demonstrated high potential for determining the fate of biogenic nutrient inputs occurring in a complex matrix such as soil. It opens up considerable opportunities to link the biological traits of specific soil engineers with geochemical processes operating at the microbial scale in the spatial structure created by earthworms.     

Multielement Isotope Ratios of Vegetables from Integrated and Organic Production

In this paper we discuss the use of isotope ratios as indicators of organic production. Few studies have investigated the influence of plant nutrition on the isotopic signatures of plants. As plant nutrition is often significantly different between integrated and organic production systems the isotope ratios in the plants may reflect this. Plant samples from a 2-year field-experiment were analyzed for ¹⁵N, ¹³C and ³⁴S content of the bulk-material and ¹⁸O-content of the leaf water. In this experiment cabbages (Brassica oleracea v. capitata f. alba cv. Rolly), onions (Allium cepa cv. Alisa Craig), lettuces (Lactuca sativa v. capitata cv. Ponchito) and Chinese cabbage (Brassica pekinesis cv. Parkin) were cultivated according to good agricultural practices for integrated and organic production. No differences in the δ ³⁴S and δ ¹⁸O values of the plants grown under the two production systems were observed. The organically produced vegetables were significantly enriched in ¹⁵N and depleted in ¹³C compared to those grown under the integrated system.     

Natural 15N abundance of epiphytes depends on the position within the forest canopy: source signals and isotope fractionation

The natural ¹⁵N abundance (δ¹⁵N) of epiphytes and its N sources were studied in the canopy of a lowland rainforest in Costa Rica. Vascular and non‐vascular epiphytes and canopy soils were collected from four canopy zones and analysed for N contents and δ¹⁵N signals. In addition, the N concentrations and δ¹⁵N signatures of bulk precipitation, throughfall and stemflow were measured during the wet and the dry season. The δ¹⁵N values of epiphyte leaves decreased significantly from the lower zones (means of ?3·9 and ?4·3‰) to the upper zones (means of ?5·4 and ?6·1‰) of the canopy. In contrast, δ¹⁵N signatures of canopy soils (average ?0·3‰) differed little between the zones. Bulk deposition was enriched in ¹⁵N (+4·3‰) compared to all other potential N sources and was higher than throughfall and stemflow (+0·5 to ?1·3‰). δ¹⁵N values of atmospheric deposition were inversely related to those of the epiphyte leaves, whereas N isotopic composition of canopy soils did not vary significantly. Consequently, it is concluded that the variations in foliar N isotope composition of epiphytes were not simply caused by utilization of isotopically different N sources, but by different ¹⁵N discrimination during N acquisition.     

A15N tracer study to compare nitrogen supply by Azolla and ammonium sulphate to IR8 rice plants grown under flooded conditions

Summary A pot experiment was carried out using a Bangladesh sandy loam paddy soil of pH 6.9 to compare the rates at which nitrogen from Azolla and ammonium sulphate was available to a high yielding rice variety, IR8, grown for 60 days in pots with 4 cm standing flood water.¹⁵N tracer studies confirm that nitrogen from ammonium sulphate was more available to the rice plants than from Azolla. An application of 6, 9 and 18 mg N of Azolla pot^–1 (each pot contained 250 g soil) increased shoot dry matter yields by 13, 29 and 49% for an uptake of 19, 36 and 85% more nitrogen; the corresponding increases on using ammonium sulphate were 33, 54 and 114% for an increased uptake of 57, 90 and 177% more nitrogen, respectively. About 34% of applied¹⁵N of Azolla was taken up by the rice plants in 60 days but 61% of¹⁵N of the ammonium sulphate was absorbed during this period. About 45% of the Azolla-N was released in 60 days, 55% remained in the soils as undecomposed material and 11% was lost as gas. The gaseous loss of¹⁵N from ammonium sulphate was 14%; 25% remained in the soils.     

Sulfur isotope inventories of atmospheric deposition, spruce forest floor and living Sphagnum along a NW–SE transect across Europe

At five European sites, differing in atmospheric Sinputs by a factor of 6, and differing in S isotope signatures ofthese inputs by up to 14 (CDT), we investigated thedirection and magnitude of an assimilation-related ³⁴S shiftand the relationship between atmospheric deposition and Sretention in selected ecosystem compartments. Bulk precipitationand spruce throughfall were collected between 1994 and 1996 inthe Isle of Mull (Scotland), Connemara (Ireland), Thorne Moors(England), Rybárenská slat' and Oceán (both Czech Republic) andanalyzed for sulfate concentrations and ³⁴S ratios. Eighteenreplicate samples per site of living Sphagnum collected inunforested peatlands and 18 samples of spruce forest floorcollected near each of the peatlands were also analyzed for Sconcentrations and ³⁴S ratios. Assimilation of S was associatedwith a negative ³⁴S shift. Plant tissues systematicallypreferred the light isotope ³²S, on average by 2. There wasa strong positive correlation between the non-marine portion ofthe atmospheric S input and total S concentration in forest floorand Sphagnum, respectively (R = 0.97 and R = 0.85). Elevated Sinputs lead to higher S retention in these two organic-richcompartments of the ecosystem. It follows that equal emphasismust be placed on organic S as on adsorption/desorption ofinorganic sulfate when studying acidification reversal inecosystems. The sea-shore sites had rainfall enriched in theheavy isotope ³⁴S due to an admixture of sea-spray. The inlandsites had low ³⁴S reflecting ³⁴S of sulfur emitted from localcoal-burning power stations. Sphagnum had always lower S contentsand higher ³⁴S ratios compared to forest floor. The within-siterange of ³⁴S ratios of Sphagnum and forest floor was wide (upto 12) suggesting that at least six replicate samples shouldbe taken when using ³⁴S as a tracer.     

Effects of inorganic nitrogen and phosphorus supply on nitrogen aquisition from alanine by birch seedlings in symbiosis with Paxillus involutus

Northern forests are exposed to relatively high ammonia inputs due to high atmospheric deposition and the common practise of forest fertilization. It is not known how increased soil NH₄ ⁺concentrations affect acquisition of symbiosis-mediated N from organic sources. We examined the effect of inorganic N and P availability on N acquisition from alanine by 43 weeks old birch (Betula pendula) seedlings in symbiosis with the ectomycorrhizal fungus Paxillus involutus. The seedlings were exposed for 9 weeks to nutrient additions equivalent to 43 kg N and 6.4 kg P ha^-1 (low N and P availability), 250 kg N and 38 kg P ha^-1(high N and P availability) or to 250 kg N and 6.4 kg P ha^-1 (high N and low P availability). Carbon and nitrogen allocation between the symbionts was assessed by exposing the foliage to ¹⁴CO₂ and the mycelium to ¹⁵N-alanine or ¹⁵NH₄ ⁺ simultaneously and measuring the distribution of the isotopic tracers after a three-day chase period. High inorganic N combined with low P availability did not have marked effect on symbiosis-mediated N uptake from alanine, whilst high N and P availability reduced alanine-derived ¹⁵N translocation by the fungus to the plant. Shoot ¹⁵N concentration and concentration of ¹⁴C in the extramatrical mycelium correlated significantly across treatments pointing to controlled reciprocity of transactions between the partners.     

Shifts in the trophic base of intermittent stream food webs

Understanding spatial and temporal variation in the trophic base of stream food webs is critical for predicting population and community stability, and ecosystem function. We used stable isotope ratios (¹³C/¹²C, and ¹⁵N/¹⁴N) to characterize the trophic base of two streams in the Ozark Mountains of northwest Arkansas, U.S.A. We predicted that autochthonous resources would be more important during the spring and summer and allochthonous resources would be more important in the winter due to increased detritus inputs from the riparian zone during autumn leaf drop. We predicted that stream communities would demonstrate increased reliance on autochthonous resources at sites with larger watersheds and greater canopy openness. The study was conducted at three low-order sites in the Mulberry River Drainage (watershed area range: 81–232 km²) seasonally in 2006 and 2007. We used circular statistics to examine community-wide shifts in isotope space among fish and invertebrate consumers in relation to basal resources, including detritus and periphyton. Mixing models were used to quantify the relative contribution of autochthonous and allochthonous energy sources to individual invertebrate consumers. Significant isotopic shifts occurred but results varied by season and site indicating substantial variation in the trophic base of stream food webs. In terms of temporal variation, consumers shifted toward periphyton in the summer during periods of low discharge, but results varied during the interval between summer and winter. Our results did not demonstrate increased reliance on periphyton with increasing watershed area or canopy openness, and detritus was important at all the sites. In our study, riffle–pool geomorphology likely disrupted the expected spatial pattern and stream drying likely impacted the availability and distribution of basal resources.     

Insights into the spatiotemporal component of feeding ecology: an isotopic approach for conservation management sciences

Aim Bringing new approaches to trace spatiotemporal variations in animals’ feeding ecology, which is fundamental for wildlife management and conservation since the accessibility of animals to trophic resources plays a key role in the dynamics of populations and metapopulations. Location Western Mediterranean coast. Methods The analysis of naturally occurring stable isotopes constitutes an exceptional approach to assess variations in the trophic ecology of species within the spatiotemporal dimensions. Here, we examined the spatiotemporal heterogeneity in resource exploitation of a nuisance and overpopulated gull species with a great feeding plasticity, the Yellow‐legged gull Larus michahellis, by measuring the stable isotope ratios of carbon (¹³C/¹²C, δ¹³C), nitrogen (¹⁵N/¹⁴N, δ¹⁵N) and sulphur (³⁴S/³³S, δ³⁴S) in different wing feathers. Results Tracing isotopic changes among feathers in relation to moulting sequence, we showed that isotopic analyses on first primary feathers were good indicators of breeding trophic ecology, while others, such as eighth secondary feathers, reflected the feeding behaviour during the non‐breeding period. This knowledge was then used to report on estimations of seasonal feeding patterns throughout the year for seven gull populations along the Western Mediterranean coast. Main conclusions The high diversity in the exploitation patterns of the foraging habitats found in the study area both at spatial and at temporal scale indicated the opportunistic feeding behaviour of Yellow‐legged gulls. Proposed isotopic approach has revealed as a useful tool to evaluate spatiotemporal variations in the feeding ecology of populations which may become clue for dealing with the management of wild species, not only nuisance populations but also endangered species. We finally pointed out the relevance of the isotopic variability among individuals to infer diet diversity and food availability of a given population, thereby allowing demographic forecasts when trophic resources vary in abundance.     

STABLE ISOTOPE ANALYSES OF TOOTH ANNULI REVEAL TEMPORAL DIETARY RECORDS: AN EXAMPLE USING STELLER SEA LIONS

Stable isotope analysis of teeth of marine mammals can provide valuable information on trophic level and source of feeding. However, the isotopic analysis of whole teeth presents only an average dietary estimate for individuals across the period of growth of that tooth. While such analyses can be valuable, particularly in the case of fossil material, in contrast, isotopic analysis of individual annuli of teeth can provide dietary information for each year of tooth growth, in some cases representing the whole of the animal's life. We measured stable-carbon isotope ratios (¹³C/¹²C) in the inorganic (hydroxyapatite) and stable-nitrogen isotope ratios (¹⁵N/¹⁴N) in the organic (primarily collagenous) components of individual tooth annuli of 18 male Steller sea lions (Eumetopias jubatus) obtained from archived collections from the Bering Sea and Gulf of Alaska and from single northern fur seals (Callorbinus ursinus) and northern elephant seals (Mirounga angustirostris) from the central Aleutian Islands and eastern Gulf of Alaska, respectively. In several individuals, we detected considerable variation in stable isotope values among annuli, up to 6.1%_O for δ¹⁵N and 5.1%_O for δ¹³C values. Enrichment in δ¹⁵N and depletion of δ¹³C values in the first annulus may correspond to dietary inputs from mother's milk during the period of suckling. Other variations among years may be caused by dietary changes or movements of individuals between regions differing in isotopic signatures of foodweb primary production. Our study indicates that the isotopic signatures of foodweb primary production. Our study indicates that the isotopic analysis of individual tooth annuli represents a fine-seale tool for dietary reconstructions involving marine mammals, and cautions against the use of whole-tooth material averaged over several annuli.