Reliability of δ13C and δ15N in faeces for reconstructing savanna herbivore diet

We tested the reliability of herbivore faecal δ¹³C and δ¹⁵N values for reconstructing diet through review of an extensive database derived from a 3-year study of ungulates in South Africa's Kruger National Park. Faeces are a useful material for stable isotope studies of diet because they record dietary turnover at very short time scales, and because sampling is non-invasive. However, the validity of faecal isotope proxies may be questioned because they represent only undigested food remains. Results from Kruger Park confirm that free-ranging browsers have faecal δ¹³C consistent with C₃ feeding, grazer faeces are C₄, and mixed-feeder faeces intermediate. Although the respective ranges do not overlap, there is significant variation in faecal δ¹³C of browsers and grazers (～2.0–4.0‰) across space and through time. We demonstrate that most (～70%) of this variation can be ascribed to corresponding patterns of variation in the δ¹³C of C₃ and C₄ plants, respectively, re-enforcing the fidelity of faecal isotope proxies for diet but highlighting a need for mixing models that control for variations in plant δ¹³C in order to achieve accurate diet reconstructions. Predictions for the effects of climate (rainfall) and ecophysiology on ¹⁵N-abundance variations in mammals do not persist in faeces. Rather, faecal δ¹⁵N tracks changes in plant δ¹⁵N, with further fractionation occurring primarily due to variations in dietary protein (reflected by %N). Controlling for these effects, we show that a dual-isotope multiple source mixing model (Isosource) can extend diet reconstructions for African savanna herbivores beyond simplified C₃/C₄ distinctions, although further understanding of variations in mammal δ¹⁵N are needed for greater confidence in this approach.     

Patterns of foliar δ15N and their control in Eastern Asian forests

Foliar δ¹⁵N has been used increasingly in research on ecosystem nitrogen (N) cycling, because it can serve as an integrator of ecosystem N cycling and thus has a potential to reveal temporal and spatial patterns of N cycling as well as how the N cycle is altered by disturbances. However, the current understanding on controls of foliar δ¹⁵N is based principally on studies from America, Europe, Australia and Africa. Here we compiled data from 65 forests at 33 sites across East Asia to explore regional patterns and what controls foliar δ¹⁵N by linking it to climate, species composition, soil depth, slope position, N deposition, and soil N availability. In East Asia, foliar δ¹⁵N ranged from ?7.1 to +2.7‰. Mean foliar δ¹⁵N values for tropical, subtropical and temperate forests were all ?3.1‰, which was unexpected. The patterns of foliar δ¹⁵N with precipitation, temperature and altitude were not clear. The variation in foliar δ¹⁵N among species and between different slope positions appeared to be small within a given forest. The δ¹⁵N for both bulk soil N and extractable inorganic N generally increased with soil depth as expected, strengthening the idea that deep-rooted trees may have access to ¹⁵N-enriched N. Different from the positive correlations reported across America and Europe, in East Asia we found that foliar δ¹⁵N decreased with increasing N deposition and did not relate to soil N availability. These discrepancies deserve more research to elucidate the mechanisms by which foliar δ¹⁵N is affected by ecosystem N availability at a regional scale.     

Variation in hair δ13C and δ15N values in long-tailed macaques (Macaca fascicularis) from Singapore

Much of the primatology literature on stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) has focused on African and New World species, with comparatively little research published on Asian primates. Here we present hair δ¹³C and δ¹⁵N isotope values for a sample of 33 long-tailed macaques from Singapore. We evaluate the suggestion by a previous researcher that forest degradation and biodiversity loss in Singapore have led to a decline in macaque trophic level. The results of our analysis indicated significant spatial variability in δ¹³C but not δ¹⁵N. The range of variation in δ¹³C was consistent with a diet based on C₃ resources, with one group exhibiting low values consistent with a closed canopy environment. Relative to other macaque species from Europe and Asia, the macaques from Singapore exhibited a low mean δ¹³C value but mid-range mean δ¹⁵N value. Previous research suggesting a decline in macaque trophic level is not supported by the results of our study.     

Soil nitrification and foliar δ15N declined with stand age in trembling aspen and jack pine forests in northern Alberta,Canada

Aims

Seed germination and seedling emergence are vulnerable to water stress in arid environments. When precipitation is low and unpredictable during the early growing season, seeds near the sand surface often suffer from hydration/dehydration during germination. We investigated the responses of seedling emergence and survival of a sand dune grass with high sand stabilization value to amount and frequency of precipitation and depth of burial in sand.

Methods

Effects of amount and frequency of precipitation, burial and hydration/dehydration on seedling emergence of Leymus secalinus, were examined using standard procedures.

Results

Seedling emergence was affected by amount and frequency of monthly precipitation and depth of burial, and it decreased as precipitation frequency decreased with same amount of precipitation. Highest emergence percentage was obtained with 100 or 150 mm precipitation at 1–4 cm depth. Hydration/dehydration treatments decreased germination and increased dormancy percentage. Young seedlings with root lengths of 0–1 mm desiccated up to 30 days revived after rehydration.

Conclusions

Seedling emergence of L. secalinus is adapted to 150 mm monthly precipitation with frequency of 10–30 times per month, 1–4 cm burial depth and dehydration interval of 1–2 days. Alteration of amount and/or frequency of precipitation caused by climate change could markedly affect seedling emergence and population regeneration of this species.     

White spruce foliar δ13C and δ15N indicate changed soil N availability by understory removal and N fertilization in a 13-year-old boreal plantation

Background and Aims

Phosphorus (P) mineralisation from crop residues is usually predicted from total P or carbon: phosphorus (C: P) ratios. However, these measures have limited accuracy as they do not take into account the presence of different P species that may be mineralised at different rates. In this study P speciation was determined using solution ³¹P nuclear magnetic resonance (NMR) spectroscopy to understand the potential fate of residue P in soils.

Methods

Mature above-ground biomass of eight different crops sampled from the field was portioned into stem, chaff and seed.

Results

The main forms of P detected in stem and chaff were orthophosphate (25–75 %), phospholipids (10–40 %) and RNA (5–30 %). Phytate was the dominant P species in seeds, and constituted up to 45 % of total P in chaff but was only detected in minor amounts (<1 %) in stem residue. The majority (65–95 %) of P in stems was water-extractable, and most of this was detected as orthophosphate. However, this includes organic P that may have been hydrolysed during the water extraction.

Conclusions

This study indicates that the majority of residue P in aboveground plant residues has the potential to be delivered to soil in a form readily available to plants and soil microorganisms.

     

Temporal records of δ13C and δ15N in North Pacific pinnipeds: inferences regarding environmental change and diet

Sea lion and seal populations in Alaskan waters underwent various degrees of decline during the latter half of the twentieth century and the cause(s) for the declines remain uncertain. The stable carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) isotope ratios in bone collagen from wild Steller sea lions (Eumetopias jubatus), northern fur seals (Callorhinus ursinus) and harbor seals (Phoca vitulina) from the Bering Sea and Gulf of Alaska were measured for the period 1951-1997 to test the hypothesis that a change in trophic level may have occurred during this interval and contributed to the population declines. A significant change in '¹⁵N in pinniped tissues over time would imply a marked change in trophic level. No significant change in bone collagen '¹⁵N was found for any of the three species during the past 47 years in either the Bering Sea or the Gulf of Alaska. However, the ¹⁵N in the Steller sea lion collagen was significantly higher than both northern fur seals and harbor seals. A significant decline in '¹³C (almost 2 ‰ over the 47 years) was evident in Steller sea lions, while a declining trend, though not significant, was evident in harbor seals and northern fur seals. Changes in foraging location, in combination with a trophic shift, may offer one possible explanation. Nevertheless, a decrease in '¹³C over time with no accompanying change in '¹⁵N suggests an environmental change affecting the base of the foodweb rather than a trophic level change due to prey switching. A decline in the seasonal primary production in the region, possibly resulting from decreased phytoplankton growth rates, would exhibit itself as a decline in '¹³C. Declining production could be an indication of a reduced carrying capacity in the North Pacific Ocean. Sufficient quantities of optimal prey species may have fallen below threshold sustaining densities for these pinnipeds, particularly for yearlings and subadults who have not yet developed adequate foraging skills.     

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.     

Vegetation recovery alters soil N status in subtropical karst plateau area: Evidence from natural abundance δ15N and δ18O

Plant and Soil - Forest recovery from disturbance can alter soil nitrogen (N) status as a result of complex interactions in plant-soil system. The δ15N and δ18O are indicators that...     

Differences in δ15N and δ13C between embryonic and maternal tissues of the ovoviviparous bluntnose sixgill shark Hexanchus griseus

Stable nitrogen (δ¹⁵N) and carbon (δ¹³C) isotope ratios from muscle, liver and yolk were analysed from the mother and embryos of an ovoviviparous shark, Hexanchus griseus. Embryonic liver and muscle had similar δ¹⁵N and δ¹³C ratios or were depleted in heavy isotopes, compared to the same maternal somatic and reproductive yolk tissues, but no relationship existed between δ¹⁵N or δ¹³C and embryo length, as expected, because a switch to placental nourishment is lacking in this species. This study expands the understanding of maternal nourishment and embryonic stable isotope differences in ovoviviparous sharks.     

Food web structure in the high Arctic Canada Basin: evidence from δ13C and δ15N analysis

The food-web structure of the Arctic deep Canada Basin was investigated in summer 2002 using carbon and nitrogen stable isotope tracers. Overall food-web length of the range of organisms sampled occupied four trophic levels, based on 3.8 trophic level enrichment (¹⁵N range: 5.3–17.7). It was, thus, 0.5–1 trophic levels longer than food webs in both Arctic shelf and temperate deep-sea systems. The food sources, pelagic particulate organic matter (POM) (¹³C=–25.8, ¹⁵N=5.3) and ice POM (¹³C=–26.9, ¹⁵N=4.1), were not significantly different. Organisms of all habitats, ice-associated, pelagic and benthic, covered a large range of ¹⁵N values. In general, ice-associated crustaceans (¹⁵N range 4.6–12.4, mean 6.9) and pelagic species (¹⁵N range 5.9–16.5, mean 11.5) were depleted relative to benthic invertebrates (¹⁵N range 4.6–17.7, mean 13.2). The predominantly herbivorous and predatory sympagic and pelagic species constitute a shorter food chain that is based on fresh material produced in the water column. Many benthic invertebrates were deposit feeders, relying on largely refractory material. However, sufficient fresh phytodetritus appeared to arrive at the seafloor to support some benthic suspension and surface deposit feeders on a low trophic level (e.g., crinoids, cumaceans). The enriched signatures of benthic deposit feeders and predators may be a consequence of low primary production in the high Arctic and the subsequent high degree of reworking of organic material.     

Ecosystem structure and soil-surface conditions drive the variability in the foliar δ13C and δ15N of Stipa tenacissima in semiarid Mediterranean steppes

We evaluated the effects of ecosystem composition and structure (species richness and diversity, cover and spatial attributes of vegetation), abiotic factors (climate and topographical features) and the condition of the bare-ground areas (evaluated using soil-surface indicators) on the performance of Stipa tenacissima [evaluated using foliar ¹³C, ¹⁵N, nitrogen concentration and the carbon-to-nitrogen (C:N) ratio] in 15 steppes of SE Spain. Foliar ¹³C values of S. tenacissima showed a low degree of variation in the studied steppes, with average values ranging from –24.1 to –22.9. Higher variation was found in the ¹⁵N values, which ranged from –5.5 to –2.4. The nitrogen concentration and the C:N ratio varied between 5.0 and 8.0 mg g^–1, and between 55.4 and 85.3, respectively. The ¹³C values became less negative with increasing spatial aggregation of perennial vegetation, while the C:N values increased with increasing perennial vegetation cover. The ¹⁵N values became more negative with increasing infiltration in the bare-ground areas, but the nitrogen concentration was not related to any of the environmental variables measured. Our results suggest that the relative importance of ecosystem structure and soil-surface conditions in the bare ground areas was higher than that of abiotic factors as determinants of the performance of S. tenacissima. The results also show that even subtle changes in these ecosystem features may lead to modifications in plant performance in semiarid S. tenacissima steppes, and thus to modifications in the associated ecosystem functions in the mid- to long-term.     

Feeding level and individual metabolic rate affect δ13C and δ15N values in carp: implications for food web studies

Stable isotope analyses are often used to calculate relative contributions of multiple food sources in an animals diet. One prerequisite for a precise calculation is the determination of the diet-tissue fractionation factor. Isotopic ratios in animals are not only affected by the composition of the diet, but also by the amount of food consumed. Previous findings regarding the latter point are controversial. As stable isotope analyses have often been used to investigate aquatic food webs, an experiment with carp (Cyprinus carpio L.) was carried out to test the influence of the feeding level and individual metabolic rate on ¹³C and ¹⁵N values of the whole body. After an initial phase, 49 carp were assigned randomly to four groups and fed the same diet at different levels for 8 weeks. For 15 fish, the energy budget was determined by indirect calorimetry. Feed and individual fish were analysed for their proximate composition, gross energy content and ¹³C and ¹⁵N values. ¹³C and ¹⁵N values differed significantly at different feeding levels. While ¹³C values of the lipids and ¹⁵N values decreased with increasing feeding rate, ¹³C values of the lipid-free matter showed a non-linear pattern. Data obtained from fish held in the respirometric system revealed a relationship between ¹³C values and the percentage retention of metabolizable energy. Our results show that reconstructing the diets of fish from the isotopic ratios when the feeding level and individual metabolic rates are unknown would introduce an error into the data used for back-calculation of up to 1 for both ¹³C and ¹⁵N values and may have substantial effects on the results of calculated diets. As other workers have pointed out, the development and application of stable isotopes to nutritional ecology studies is a field in its infancy and gives rise to erroneous, misleading results without nutritional, physiological and ecological knowledge.     

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.     

Variability in δ13C and δ15N trophic discrimination factors for teleost fishes: a meta-analysis of temperature and dietary effects

Reviews in Fish Biology and Fisheries - Stable isotope analysis (SIA) is widely used to assess animal diet and movements, requiring accurate estimates of trophic discrimination factors (TDFs)....     

Experimentally Derived δ13C and δ15N Discrimination Factors for Gray Wolves and the Impact of Prior Information in Bayesian Mixing Models

Stable isotope analysis of diet has become a common tool in conservation research. However, the multiple sources of uncertainty inherent in this analysis framework involve consequences that have not been thoroughly addressed. Uncertainty arises from the choice of trophic discrimination factors, and for Bayesian stable isotope mixing models (SIMMs), the specification of prior information; the combined effect of these aspects has not been explicitly tested. We used a captive feeding study of gray wolves (Canis lupus) to determine the first experimentally-derived trophic discrimination factors of C and N for this large carnivore of broad conservation interest. Using the estimated diet in our controlled system and data from a published study on wild wolves and their prey in Montana, USA, we then investigated the simultaneous effect of discrimination factors and prior information on diet reconstruction with Bayesian SIMMs. Discrimination factors for gray wolves and their prey were 1.97‰ for δ¹³C and 3.04‰ for δ¹⁵N. Specifying wolf discrimination factors, as opposed to the commonly used red fox (Vulpes vulpes) factors, made little practical difference to estimates of wolf diet, but prior information had a strong effect on bias, precision, and accuracy of posterior estimates. Without specifying prior information in our Bayesian SIMM, it was not possible to produce SIMM posteriors statistically similar to the estimated diet in our controlled study or the diet of wild wolves. Our study demonstrates the critical effect of prior information on estimates of animal diets using Bayesian SIMMs, and suggests species-specific trophic discrimination factors are of secondary importance. When using stable isotope analysis to inform conservation decisions researchers should understand the limits of their data. It may be difficult to obtain useful information from SIMMs if informative priors are omitted and species-specific discrimination factors are unavailable.     

Developmental and circadian control of the capacity for δ-aminolevulinic acid synthesis in green barley

The synthesis of δ-aminolevulinic acid (δ-ALA) is a key step in the regulation of tetrapyrrole synthesis. To study the developmentally and circadian-clock controlled mechanism that co-ordinates synthesis of chlorophylls and chlorophyll-binding proteins, δ-ALA-synthesising capacity was analysed in barley (Hordeum vulgare L.) primary leaves grown under dark/light or constant light conditions. The δ-ALA-forming activity oscillated within 24 h with a maximum at the transition of dark to light and a minimum 12 h later, indicating the involvement of the circadian oscillator during development. The capacity for δ-ALA synthesis increased transiently in the middle of barley primary leaves. The δ-ALA-forming-activity correlated well with the previously published steady-state level of mRNA for light-harvesting chlorophyll-binding proteins in space and time; this supports the view of a co-ordinate synthesis of chlorophyll and pigment-binding proteins. Steady-state levels of mRNAs encoding the three enzymes of the δ-ALA-synthesising pathway and of proteins for glutamyl-tRNA reductase (GluTR) and glutamate 1-semialdehyde aminotransferase (GSA AT; EC 5.4.3.8) were analysed for their developmental and circadian expression in barley leaves. The contents of GluTR mRNA and protein cycled parallel to the changes in δ-ALA-forming activity. The levels of GSA AT mRNA oscillated in an opposite phase, but the protein content did not show substantial oscillation under diurnal and circadian growth conditions. No circadian oscillation was detected for glutamyl tRNA synthase (GluRS; EC 6.1.1.17). Maximal GluTR mRNA content and protein was observed in the middle (segments 3 and 4) of the barley primary leaves. The developmentally controlled expression of GluTR therefore differs from that of GSA AT and GluRS, but resembles the capacity for δ-ALA synthesis in a barley leaf gradient. These data indicate that the oscillating, light-dependent and spatial expression of GluTR mRNA might contribute to the regulated formation of the chlorophyll precursor δ-ALA. Received: 29 April 1996 / Accepted 11 December 1996     

δ13C and δ15N indicators of fish and shrimp community diet and trophic structure in a mangrove ecosystem in Thailand

Stable carbon and nitrogen isotope ratios were used to elucidate primary carbon sources and trophic relationships of the fish and shrimp community in the Klong Ngao mangrove ecosystem, southern Thailand. There were no significant differences in isotopic compositions of biota between mangrove and offshore sites (Welch–Aspin test). The δ¹⁵N values of eight fish species and two shrimp species at both sites were also not significantly different by the test, meaning that at both sites they feed on the same diets due to the discharge of large quantities of mangrove sediments. The δ¹⁵N isotopic enrichment of consumers suggested that there are four trophic levels in the Klong Ngao food web, with at least two fish species capable of switching feeding strategies and thus altering their apparent trophic positions. Phytoplankton culture experiments indicated that mangrove-derived sediments could play an important role in stimulating phytoplankton growth for low turbidity offshore areas, thus providing an alternate food source. The isotopic associations among sources and consumers indicated that mangroves were the major carbon source supporting aquatic food webs in the Klong Ngao ecosystem.     

Effect of the fungal endophyte Epichloë bromicola on polyamines in wild barley (Hordeum brevisubulatum) under salt stress

Plant and Soil - The endophytic fungus Epichloë bromicola forms mutualistic symbiotic associations with wild barley (Hordeum brevisubulatum) in the saline-alkali areas of northwestern China....