The trophic structure of bark-living oribatid mite communities analysed with stable isotopes (15N, 13C) indicates strong niche differentiation

The aim of the present study was to identify food sources of bark-living oribatid mites to investigate if trophic niche differentiation contributes to the diversity of bark living Oribatida. We measured the natural variation in stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) in oribatid mites from the bark of oak (Quercus robur), beech (Fagus sylvatica), spruce (Picea abies) and pine (Pinus sylvestris) trees and their potential food sources, i.e., the covering vegetation of the bark (bryophytes, lichens, algae, fungi). As a baseline for calibration the stable isotope signatures of the bark of the four tree species were measured and set to zero. Oribatid mite stable isotope ratios spanned over a range of about 13 δ units for ¹⁵N and about 7 δ units for ¹³C suggesting that they span over about three trophic levels. Different stable isotope signatures indicate that bark living oribatid mites feed on different food sources, i.e., occupy distinct trophic niches. After calibration stable isotope signatures of respective oribatid mite species of the four tree species were similar indicating close association of oribatid mites with the corticolous cover as food source. Overall, the results support the hypothesis that trophic niche differentiation of bark living oribatid mites contributes to the high diversity of the group.     

The importance of quantifying inherent variability when interpreting stable isotope field data

Stable isotope data are often used to assess diet, trophic level, trophic niche width and the extent of omnivory. Notwithstanding ongoing discussions about the value of these approaches, variations in isotopic signatures among individuals depend on inherent variability as well as differences in feeding habitats. Remarkably, the relative contributions of diet variation and inherent variability to differences in δ¹⁵N and δ¹³C among individuals have not been quantified for the same species at the same life history stages, and inherent variability has been ignored or assumed. We quantified inherent variability in δ¹³C and δ¹⁵N among individuals of a marine fish (the European sea bass, Dicentrarchus labrax) reared in a controlled environment on a diet of constant isotopic composition and compared it with variability in δ¹³C and δ¹⁵N among individuals from wild bass populations. The analysis showed that inherent variability among reared individuals on a controlled diet was equivalent to a large proportion of the observed variability among wild individuals and, therefore, that inherent variability should be measured to establish baseline variability in wild populations before any assumptions are made about the influence of diet. Given that inherent variability is known to be dependent on species, life history stage and the environment, our results show that it should be quantified on a case-by-case basis if diet studies are intended to provide absolute assessments of dietary habits.     

Diet differences among age classes of Arctic seals: evidence from stable isotope and mercury biomarkers

A basic understanding of current food web dynamics and baseline data from which to measure future change is necessary to understand species re-distribution and altered competition for food with climate change. We use mercury (Hg) and carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope ratios as biomarkers to understand species diet differences and age class differences among ringed (Phoca hispida), bearded (Erignathus barbatus), and harbour (P. vitulina) seals in a subarctic marine ecosystem. Adult bearded seals had significantly lower δ¹⁵N and muscle Hg than bearded seal pups, whereas the opposite was observed in ringed seals where pups had lower δ¹⁵N than adults, suggesting age specific foraging differences in trophic food level for both species. For harbour seals, δ¹⁵N did not differ significantly among ages while Hg and δ¹³C did. The δ¹³C in muscle supports that bearded seals in this study are benthic feeders and are part of a separate food web from ringed seals and harbour seals. Harbour seals had the highest levels of mercury and δ¹⁵N, indicating they feed at a higher trophic level relative to the other two seal species. Carbon and nitrogen isotopic ratios and Hg levels illustrate how resources are partitioned among three seal species and offer evidence for separation based on life stages within species.     

Stable carbon and nitrogen isotope ratio profiling of sperm whale teeth reveals ontogenetic movements and trophic ecology

Teeth from male sperm whales (Physeter macrocephalus) stranded in the North-eastern Atlantic were used to determine whether chronological profiles of stable isotope ratios of C (δ¹³C) and N (δ¹⁵N) across dentine growth layers could be used to detect known ontogenetic benchmarks in movements and trophic ecology. Profiles showed a general decrease in δ¹³C (median = 1.91‰) and an increase in δ¹⁵N (median = 2.42‰) with age. A marked decline in δ¹³C occurred for all 11 teeth around 9–10 years and again for six individuals around 20 years. After the early twenties the δ¹³C continued to decline with age for all teeth. These results are consistent with males segregating from natal groups in low latitudes with the onset of puberty between 4 and 15 years and gradually dispersing pole-ward into ¹³C-depleted temperate waters. Penetration into further depleted, productive high latitudes after the age of 20 might facilitate the spurt of accelerated growth rate observed around this age. Breeding migrations back to lower latitudes were not reflected in the δ¹³C profiles possibly due to being short compared to the time spent feeding in high latitudes. The timings of marked isotopic change in the δ¹⁵N profiles reflect those of the δ¹³C profiles, suggesting a link between dietary changes and movements. The observed increase in δ¹⁵N with age is likely to be caused by a trophic level increase as males grow in size, probably feeding on larger prey. An additional explanation could be that, in the higher latitudes of the North Atlantic, the main prey source is the high trophic level squid Gonatus fabricii. Also, the lower latitudes from where males disperse are depleted in basal ¹⁵N. Profiles of δ¹³C and δ¹⁵N in sperm whale teeth gathered from different regions, sexes, and periods in time, could provide a unique way to understand the ecology of this species across different oceans.     

Food web structure of a shallow eutrophic lake (Lake Taihu,China) assessed by stable isotope analysis

Lake Taihu is a large, shallow, and eutrophic lake in China. It has provided local communities with valuable fisheries for centuries, but little is known of the trophodynamics, or of its faunal communities. Carbon and nitrogen isotopic composition was used to assess its trophic pathways and the food web structure [food sources and trophic levels (TL)]. Basal food sources were distinguishable based on their δ¹³C values, ranging from −27.2 to −15.2‰. Consumers were also well separated in δ¹³C (−26.9 to −17.9‰ for invertebrates and −25.7 to −18.1‰ for fishes), which allowed for an effective discrimination of carbon sources between these fauna. An average trophic enrichment factor of 3.4‰ was used to calculate the TLs based on δ¹⁵N of zooplankton, with results indicating a food web having four TLs. Although δ¹⁵N values overlap and cover a large range within trophic compartments, the isotopic signatures of the species assessed revealed a general trend of ¹⁵N enrichment with increasing TL. Stable isotope signatures were also used to establish a general food web scheme in which five main trophic pathways were analyzed.     

Temporal stability of niche use exposes sympatric Arctic charr to alternative selection pressures

There is now strong evidence that foraging niche specialisation plays a critical role in the very early stages of resource driven speciation. Here we test critical elements of models defining this process using a known polymorphic population of Arctic charr from subarctic Norway. We test the long-term stability of niche specialisation amongst foraging predators and discuss the possibility that contrasting foraging specialists are exposed to differing selection regimes. Inter-individual foraging niche stability was measured by combining two time-integrated ecological tracers of the foraging niche (each individual’s δ¹³C and δ¹⁵N stable isotope (SI) signatures and their food borne parasite fauna) with a short-term measure of foraging niche use (stomach contents composition). Three dietary subgroups of predators were identified, including zooplankton, gammarid and benthivore specialists foragers. Zooplanktivorous specialists had muscle low in δ ¹³C, a high abundance of parasites transmitted from pelagic copepods, a smaller head, longer snout and a more slender body-form than gammaridivorous specialist individuals which had muscle more enriched in δ ¹³C and high abundance of parasites transmitted from benthic Gammarus. Benthivorous individuals were intermediate between the other two foraging groups according to muscle SI-signals (δ¹³C) and loadings of parasites transmitted from both copepods and Gammarus. The close relationship between subgroups identified by stomach contents, time-integrated tracers of niche use (SI and parasites) and functional trophic morphology (niche adaptations) demonstrate a long-term temporally stable niche use of each individual predator. Differential habitat use and contrasting parasite communities and loadings, show differential exposure to different suites of selection pressures for different foraging specialists. Results also show that individual specialisation in trophic behaviour and thus exposure to different suites of selection pressures are stable over time, and thus provide a platform for disruptive selection to operate within this sympatric system.     

Spatial and temporal variabilities of δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N within lower trophic levels of a large lake: implications for estimating trophic relationships of consumers

Stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) often have unique values among lake habitats (e.g. benthic, littoral, pelagic), providing a widely used tool for measuring the structure and energy flow in aquatic food webs. However, there has been little recognition of the spatial and temporal variabilities of these isotopes within habitats of aquatic ecosystems. To address this, δ¹³C and δ¹⁵N were measured in seston, zebra mussels (Dreissena polymorpha) and young-of-year (YOY) yellow (Perca flavescens), and white perch (Morone americana) collected from four sites across the offshore habitat of the western basin of Lake Erie during June–September 2009. Values of δ¹³C and δ¹⁵N showed significant spatial and temporal variations, with month accounting for >50% of the variation, for both stable isotopes and all the species except seston. Such variation in isotope values has the potential to significantly influence or confound interpretation of stable isotopes in measures, such as trophic position (TP) which use lower trophic level organisms as their baseline. For example, TP was found to vary up to 0.7 for yellow and white perch (TP = δ¹⁵N_fish − δ¹⁵N_{zebra mussel}/diet-tissue fractionation factor) depending on the zebra mussel data used (e.g., from a different location or a different collection month). As the use of stable isotopes continues to move from qualitative to more quantitative measures of trophic structure, food web research must recognize the importance of stable isotopes' variability in lower trophic level organisms, especially in large lake systems.     

Elucidating the trophodynamics of four coral reef fishes of the Solomon Islands using δ15N and δ13C

Size-related diet shifts are important characteristics of fish trophodynamics. Here, body size–related changes in muscle δ¹⁵N and δ¹³C of four coral reef fishes, Acanthurus nigrofuscus (herbivore), Chaetodon lunulatus (corallivore), Chromis xanthura (planktivore) and Plectropomus leopardus (piscivore) were investigated at two locations in the Solomon Islands. All four species occupied distinct isotopic niches and the concurrent δ¹³C′ values of C. xanthura and P. leopardus suggested a common planktonic production source. Size-related shifts in δ¹⁵N, and thus trophic level, were observed in C. xanthura, C. lunulatus and P. leopardus, and these trends varied between location, indicating spatial differences in trophic ecology. A literature review of tropical fishes revealed that positive δ¹⁵N-size trends are common while negative δ¹⁵N-size trends are rare. Size-δ¹⁵N trends fall into approximately equal groups representing size-based feeding within a food chain, and that associated with a basal resource shift and occurs in conjunction with changes in production source, indicated by δ¹³C. The review also revealed large scale differences in isotope-size trends and this, combined with small scale location differences noted earlier, highlights a high degree of plasticity in the reef fishes studied. This suggests that trophic size analysis of reef fishes would provide a productive avenue to identify species potentially vulnerable to reef impacts as a result of constrained trophic behaviour.     

Preliminary assessment of Greenland halibut diet in Cumberland Sound using stable isotopes

We provide preliminary carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope assessment of the Greenland halibut (Reinhardtius hippoglossoides) diet in Cumberland Sound, with focus on two possible prey sources: pelagic represented by capelin (Mallotus villosus) and epibenthic represented by shrimp (Lebbeus polaris). The δ¹³C for the Greenland halibut stock indicated a pelagic carbon source in Cumberland Sound while stable isotope mixing models, IsoSource and MixSIR, indicated a 99% dietary composition of capelin relative to the shrimp. The δ¹⁵N did not vary across Greenland halibut size ranges and placed them at a fourth trophic position relative to a primary herbivore. This study provides the starting point for more elaborate Cumberland Sound research on the local Greenland halibut feeding ecology by confirming pelagic feeding and establishing relative trophic position as well as identifying stable isotopes as a useful tool for the study of diet in cold water fish species.     

Investigation of mercury concentrations in fur of phocid seals using stable isotopes as tracers of trophic levels and geographical regions

Recent studies have shown that the complementary analysis of mercury (Hg) concentrations and stable isotopic ratios of nitrogen (δ¹⁵N) and carbon (δ¹³C) can be useful for investigating the trophic influence on the Hg exposure and accumulation in marine top predators. In this study, we propose to evaluate the interspecies variability of Hg concentrations in phocids from polar areas and to compare Hg bioaccumulation between both hemispheres. Mercury concentrations, δ¹⁵N and δ¹³C were measured in fur from 85 individuals representing 7 phocidae species, a Ross seal (Ommatophoca rossii), Weddell seals (Leptonychotes weddellii), crabeater seals (Lobodon carcinophagus), harbour seals (Phoca vitulina), grey seals (Halichoerus grypus), ringed seals (Pusa hispida) and a bearded seal (Erignathus barbatus), from Greenland, Denmark and Antarctica. Our results showed a positive correlation between Hg concentrations and δ¹⁵N values among all individuals. Seals from the Northern ecosystems displayed greater Hg concentrations, δ¹⁵N and δ¹³C values than those from the Southern waters. Those geographical differences in Hg and stable isotopes values were likely due to higher environmental Hg concentrations and somewhat greater number of steps in Arctic food webs. Moreover, dissimilarities in feeding habits among species were shown through δ¹⁵N and δ¹³C analysis, resulting in an important interspecific variation in fur Hg concentrations. A trophic segregation was observed between crabeater seals and the other species, resulting from the very specific diet of krill of this species and leading to the lowest observed Hg concentrations.     

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

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

Trophic polymorphism in bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>) introduced into Lake Biwa: evidence from stable isotope analysis

Trophic polymorphism was recently reported in introduced bluegill (Lepomis macrochirus) in Lake Biwa, Japan, where three morphs are specialized in benthic invertebrates (benthivorous type), submerged aquatic plants (herbivorous type), and zooplankton (planktivorous type). We evaluated the long-term effects of food resource utilization by these trophic morphs using stable isotope ratios, δ¹⁵N and δ¹³C. A significant difference in δ¹⁵N was found between the benthivorous and planktivorous types. The planktivorous type had the higher δ¹⁵N value, which corresponded with the value expected from its prey, zooplankton. The lower δ¹⁵N value of the benthivorous type would be derived from the lower δ¹⁵N values of benthic prey organisms compared to zooplankton. These results support previous findings that the benthivorous and planktivorous types have different food resource utilization. In contrast, the δ¹⁵N and δ¹³C values of the herbivorous type were distinctly different from the expected values, indicating that this type was unlikely to utilize aquatic plants substantially, contradicting the results of the dietary analysis.     

Isotopic signatures and trophic status of Ramaria

The genus Ramaria is composed of several subgenera that often correspond to specific trophic strategies. Because carbon and nitrogen isotopes can be used to assess fungal trophic status and nitrogen sources, we accordingly carried out an extensive survey of isotopic patterns in archived specimens of Ramaria from Germany and other locations. Isotopic patterns in species generally corresponded to subgeneric affiliations and to the range of different potential substrates, with fungi fruiting on wood and litter (subgenera Asteroramaria and Lentoramaria) much lower in δ¹⁵N (≈−3‰) than ectomycorrhizal taxa (≈12‰) (subgenus Ramaria) or taxa fruiting on soil (≈13‰) (subgenus Echinoramaria). Conversely, fungi fruiting on wood and litter were higher in δ¹³C (−23‰) than those fruiting on soil (≈−27‰), with ectomycorrhizal fungi intermediate (≈−24.5‰). Fungi colonizing mineral soil horizons were about 3‰ enriched in ¹⁵N relative to those colonizing both mineral and organic horizons. The high δ¹⁵N and low δ¹³C signatures of taxa fruiting on soil remains unexplained. The high degree of fidelity of isotopic signatures with subgeneric classifications and life history traits suggests that sporocarps are good integrators of patterns of carbon and nitrogen cycling for specific taxa. Archived specimens represent a useful trove of life history information that could be mined without requiring extensive supporting isotopic data from other ecosystem pools.     

Stable carbon and nitrogen incorporation in blood and fin tissue of the catfish <Emphasis Type="Italic">Pterygoplichthys disjunctivus</Emphasis> (Siluriformes,Loricariidae)

A feeding trial was performed in the laboratory with the catfish species Pterygoplichthys disjunctivus to determine stable carbon (¹³C) and nitrogen (¹⁵ N) turnover rates and discrimination factors in non-lethally sampled tissues (red blood cells, plasma solutes, and fin). A second feeding trial was conducted to determine what P. disjunctivus could assimilate from low-quality wood-detritus—refractory polysaccharides (e.g., cellulose), or soluble wood-degradation products inherent in wood-detritus. This was performed by feeding the fish an artificial wood-detritus diet with fibrous (δ¹³C = −26.36‰; δ¹⁵  N = 2.13‰) and soluble portions (δ¹³C = −11.82‰; δ¹⁵  N = 3.39‰) that had different isotopic signatures and monitoring the dynamics of isotopic incorporation in the different tissues over time. Plasma solutes turned over more quickly than red blood cells for ¹³C and ¹⁵ N. However, in contrast to previous studies of juvenile fishes, C and N incorporation was primarily driven by catabolic tissue turnover as opposed to growth rate. Tissue-diet discrimination factors for ¹⁵ N varied from 4.08 to 5.17‰, whereas they were <2‰ for ¹³C (and less than 0.3‰ for plasma and red blood cells). The results of trial two suggested that P. disjunctivus could not assimilate refractory polysaccharides. Moreover, the δ¹³C and δ¹⁵ N signatures of wild-caught P. disjunctivus from Florida confirmed their detrital trophic standing in Floridian aquatic ecosystems.     

Variability and directionality of temporal changes in δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N of aquatic invertebrate primary consumers

Seasonal oscillations in the carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope signatures of aquatic algae can cause seasonal enrichment–depletion cycles in the isotopic composition of planktonic invertebrates (e.g., copepods). Yet, there is growing evidence that seasonal enrichment–depletion cycles also occur in the isotope signatures of larger invertebrate consumers, taxa used to define reference points in isotope-based trophic models (e.g., trophic baselines). To evaluate the general assumption of temporal stability in non-zooplankton aquatic invertebrates, δ¹³C and δ¹⁵N time series data from the literature were analyzed for seasonality and the influence of biotic (feeding group) and abiotic (trophic state, climate regime) factors on isotope temporal patterns. The amplitude of δ¹³C and δ¹⁵N enrichment–depletion cycles was negatively related to body size, although all size-classes of invertebrates displayed a winter-to-summer enrichment in δ¹³C and depletion in δ¹⁵N. Among feeding groups, periphytic grazers were more variable and displayed larger temporal changes in δ¹³C than detritivores. For nitrogen, temporal variability and magnitude of directional change of δ¹⁵N was most strongly related to ecosystem trophic state (eutrophic > mesotrophic, oligotrophic). This study provides evidence of seasonality in the isotopic composition of aquatic invertebrates across very broad geographical and ecological gradients as well as identifying factors that are likely to modulate the strength and variability of seasonality. These results emphasize the need for researchers to recognize the likelihood of temporal changes in non-zooplankton aquatic invertebrate consumers at time scales relevant to seasonal studies and, if present, to account for temporal dynamics in isotope trophic models.     

Metabolic fractionation of stable carbon isotopes: implications of different proximate compositions for studies of the aquatic food webs using δ13C data

In two laboratory experiments using tilapia (Oreochromis niloticus) and common carp (Cyprinus carpio), we investigated the effect of lipid content in the fish carcass on the stable carbon isotope ratio (δ¹³C). In both experiments, a significant negative correlation was found between lipid content in the carcass dry matter and δ¹³C of total dry matter, but there was no influence on the δ¹³C of fat-free dry matter or lipids. As the lipid content of fish is known to vary with reproductive stage, season or nutritional state, separate analysis of fractions of the proximate composition of dry matter will lead to more reliable results than analysing the whole body. The differences in δ¹³C between diet and fish carcass (trophic shift) were different for the two species, calling for feeding trials under controlled conditions using the species and the feeds in question before applying the stable isotope tracer technique in the analysis of food webs. Received: 1 December 1997 / Accepted: 1 March 1998     

Research article: small-scale spatial variation of δ13C and δ15N isotopes in Antarctic carbon sources and consumers

Regional food web studies that fail to account for small-scale isotopic variability can lead to a mismatch between an organism’s inferred and true trophic position. Misinterpretation of trophic status may result, substantially limiting spatial and temporal comparability of food web studies. We sampled several carbon sources and consumers in a nested design to assess the variability of food web members across small spatial scales (100 s of m to several km) in regions around the Windmill Islands and Vestfold Hills in East Antarctica. For carbon sources, δ¹³C in sea ice POM was particularly variable between locations (km apart) and between sites (100 s of m apart) with replicate samples varying by up to 16‰. Macroalgae δ¹³C was less variable (replicate samples ranging up to 6.9‰ for the red alga Iridaea cordata), yet still differed between locations. Sediment POM and pelagic POM were the least variable, displaying minimal differences between locations or sites for δ¹³C and δ¹⁵N. Three out of eight consumers were significantly different between locations for δ¹³C, and five out of eight for δ¹⁵N, with the fish Trematomus bernacchii the most variable for both δ¹³C and δ¹⁵N. At smaller scales, the amphipod Paramorea walkeri showed significant variation between sites in δ¹³C but not in δ¹⁵N. We attribute small-scale variability to the dynamic physical environment for carbon sources in coastal systems and a close coupling of diet to habitat for consumers. We highlight the need to account for small-scale spatial variation in sampling designs for regional food web studies.     

Organic Matter Stable Isotope (δ13C, δ15N) Response to Historical Eutrophication of Lake Taihu, China

We explored the use of carbon and nitrogen isotope ratios (δ¹³C, δ¹⁵N) in sediment organic matter as proxy indicators of historical changes in the trophic state of Lake Taihu, the third largest freshwater lake in China. Stable isotope signatures in four sediment cores spanning the 20th century were compared with instrumental records of lake-water trophic state. The comparative study shows that, between ∼ ∼1950 and 1990 AD, the δ¹³C and δ¹⁵N of sediment organic matter throughout Lake Taihu increased along the trophic gradient from oligotrophy to eutrophy due to biological isotopic fractionation. However, in the 1990s, the trophic state of Lake Taihu diverged into two different trophic systems, a hypereutrophic western Lake Taihu dominated by blue-green algae and a mesoeutrophic eastern Lake Taihu dominated by vascular aquatic plants. During the post-1990 AD shift from mesoeutrophic to hypereutrophic state in western Lake Taihu, organic matter δ¹³C and δ¹⁵N decreased sharply in response to pronounced shifts in the aquatic ecosystem. The results indicate that ¹³C-depleted phytoplankton replaced macrophytes in western Lake Taihu. δ¹⁵N values in western Lake Taihu also decreased because of N₂ fixation by cyanobacteria in this highly productive ecosystem. By contrast, in eastern Lake Taihu, organic matter δ¹³C and δ¹⁵N values show a post-1990 AD trend towards slightly lower values, but they remain higher than the long-term average. This recent ¹³C–enrichment of organic matter indicates that periods of high productivity in the restricted eastern sub-basin of Lake Taihu limited aqueous CO₂ availability, causing a decrease in isotopic discrimination during photosynthesis. After ∼ ∼1990 AD, organic matter δ¹⁵N values for eastern Lake Taihu only dropped slightly, suggesting that the contribution of phytoplankton to the sediment organic matter increased slightly. Taken together, the results indicate that nitrogen-fixing cyanobacteria probably played a much smaller role in primary productivity in this part of eastern Lake Taihu, compared with western Lake Taihu. Despite the complexity of carbon and nitrogen cycles in lakes, the agreement between the stable isotope signatures and instrumental records for Lake Taihu suggests that δ¹³C and δ¹⁵N in sediment organic matter are capable of recording important shifts in the spatial and temporal evolution of lake-water trophic state.     

Fast carnivores and slow herbivores: differential foraging strategies among grizzly bears in the Canadian Arctic

Categorizing animal populations by diet can mask important intrapopulation variation, which is crucial to understanding a species’ trophic niche width. To test hypotheses related to intrapopulation variation in foraging or the presence of diet specialization, we conducted stable isotope analysis (δ¹³C, δ¹⁵N) on hair and claw samples from 51 grizzly bears (Ursus arctos) collected from 2003 to 2006 in the Mackenzie Delta region of the Canadian Arctic. We examined within-population differences in the foraging patterns of males and females and the relationship between trophic position (derived from δ¹⁵N measurements) and individual movement. The range of δ¹⁵N values in hair and claw (2.0–11.0‰) suggested a wide niche width and cluster analyses indicated the presence of three foraging groups within the population, ranging from near-complete herbivory to near-complete carnivory. We found no linear relationship between home range size and trophic position when the data were continuous or when grouped by foraging behavior. However, the movement rate of females increased linearly with trophic position. We used multisource dual-isotope mixing models to determine the relative contributions of seven prey sources within each foraging group for both males and females. The mean bear dietary endpoint across all foraging groups for each sex fell toward the center of the mixing polygon, which suggested relatively well-mixed diets. The primary dietary difference across foraging groups was the proportional contribution of herbaceous foods, which decreased for both males and females from 42–76 to 0–27% and 62–81 to 0–44%, respectively. Grizzlies of the Mackenzie Delta live in extremely harsh conditions and identifying within-population diet specialization has improved our understanding of varying habitat requirements within the population.     

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