Unravelling complexities in benthic food webs using a dual stable isotope (hydrogen and carbon) approach

1. Stable carbon isotope studies have been an essential component of research regarding the contribution of methane (CH₄)-derived carbon to freshwater food webs and results have suggested that benthic macroinvertebrates in billabongs, streams and lakes may be partially, and in some instances, significantly 'fuelled' by methanotrophic biomass. However, the singular use of carbon isotopes can lead to ambiguous interpretations concerning the origin of carbon, especially in systems where phototrophs are likely to be using carbon respired sources and hence show more ¹³C-depleted values.
2. These uncertainties can be further resolved by the inclusion of additional isotopic data. Stable hydrogen isotopes are being increasingly used in food web studies with a marked advantage that sources may be isotopically distinct by one or two orders of magnitude greater than stable carbon or nitrogen, the isotopes most commonly used to delineate trophic interactions. By using hydrogen as a second biogeochemical tracer we provide further supportive evidence for the assimilation of methanotrophic microbial biomass by chironomid larvae.
3. Moreover, the hydrogen and carbon isotope values we found in chironomid tissues appear to reflect the original substrate used during methanogenesis; either acetate fermentation or carbonate reduction. Use of the former tends to result in relatively heavy carbon and light hydrogen isotope values due to kinetic isotope effects, whereas use of the latter results in relatively lighter carbon and heavier hydrogen isotope values.
4. We provide preliminary evidence to suggest that hydrogen and carbon isotope values in macroinvertebrates may be used to distinguish between CH₄ formation pathways and help to explain inter-depth and inter-specific differences between co-existing chironomid species found in the same lake.     

Microwear evidence for Plio-Pleistocene bovid diets from Makapansgat Limeworks Cave, South Africa

Makapansgat Limeworks Cave is a well-known Australopithecus africanus bearing locality that has spawned a considerable amount of paleoecological research because of its hominin component. Most recently, the paleoecology of this Plio-Pleistocene site has been studied by determining the diet and habitat of other extinct taxa, particularly the bovids. The diets of seven bovids (Aepyceros sp., Gazella vanhoepeni, Makapania broomi, Parmularius braini, Redunca darti, Tragelaphus sp. aff. T. angasii, and Tragelaphus pricei) have now been classified using taxonomic uniformitarianism, ecomorphology, stable carbon isotopes, and mesowear analysis. Here, dental microwear is applied to the same bovids for additional comparison and to further elucidate the strengths and weaknesses of each method. The different dietary proxy methods noted provide a temporal continuum, with genetic signals such as ecomorphology and taxonomic uniformitarianism indicating behavioral adaptations over geologic time, while nongenetic data such as stable carbon isotopes and mesowear reflect different aspects of average diet over extended portions of an animal's life, and dental microwear provides dietary snapshots.Microwear separated an extant baseline of ten bovid species into expected dietary categories and the Makapansgat bovids clearly fell into two groups with the same degree of separation as between extant grazers and browsers. The results indicate that a multidisciplinary approach produces a more accurate and robust reconstruction of past diets. In sum, the microwear analysis is in-line with the isotope and mesowear results, which suggest a stronger browsing component than either taxonomic uniformitarianism or ecomorphology imply.     

Dietary adaptations in an ungulate community from the late Pliocene of Greece

The dietary morphological methods of mesowear and microwear were applied to ungulates of the late Pliocene fauna of Sésklo (Thessaly, Greece). The results provide evidence for the predominance of open grassland in the area, as the most common species, Equus stenonis, was a strict grazer. The rare cervid cf. Croizetoceros ramosus was the only browser. The antelopes (genera Gazella and Gazellospira) yielded discrepant microwear and mesowear results. This is interpreted as an indication of regional or seasonal dietary resource differentiation, inferring that the antelopes were probably mixed feeders that grazed occasionally or periodically.     

Contribution of Chemoautotrophic Production to Freshwater Macroinvertebrates in a Headwater Stream Using Multiple Stable Isotopes

We estimated the food sources of macroinvertebrates using carbon, nitrogen, and sulfur stable isotopes in a headwater stream. Stream food webs, including macroinvertebrates, rely on production from autochthonous and allochthonous photosynthesis. We found that a freshwater grazer, the snail Semisulcospira libertina, may assimilate different food sources, based on stable carbon and nitrogen isotope evidence from snail muscle and the much lighter sulfur isotope signature than those of other associated macroinvertebrates. Previous studies have shown the importance of methanotrophic and sulfur bacteria in reductive environments (clay and organic‐rich sediments) as food sources for macroinvertebrates. Our results show that the production by chemoautotrophic bacteria contributes to the food sources of a snail in a stream. Thus, the chemoautotrophic bacteria are important in the freshwater food webs, even in mostly aerobic habitats. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The confounding effects of source isotopic heterogeneity on consumer–diet and tissue–tissue stable isotope relationships

Stable isotope analysis of consumer tissues document patterns of resource use because data are linearly related to isotope compositions of their source(s) (i.e., food, water, etc.). Deviations in parameters estimated for these relationships can arise from variations in consumer tissue-diet spacing (Δ(TS)) and the level of isotopic heterogeneity in the source(s). We present a set of simple hypotheses that distinguish between the effects of Δ(TS) and source isotope heterogeneity. The latter may arise via mixed diets, during tissue turnover, or by isotopic routing of dietary components. We apply these concepts to stable carbon and nitrogen isotope relationships between gut contents and body tissues of large mammal herbivores from mixed C(3)/C(4) South African savannas and test predictions based on the compound- and/or time-specific data archived within each material. Predicted effects of source isotope heterogeneity are readily detected in carbon isotope relationships between materials representing different time periods or comprising bulk versus protein-only diet components. Differences in Δ(TS) of carbon isotopes across mammal herbivore species with very different feeding niches (and diet isotope compositions) are likely to be small or non-existent in these habitats. Variations in Δ(TS) estimated for nitrogen isotopes are much greater, leading to inconsistencies that cannot be explained by diet or trophic level effects alone. The effects of source heterogeneity on isotopic relationships generate numerical artefacts that have been misinterpreted as variations in Δ(TS). We caution against generalized application of hypotheses based on assumptions of source isotopic homogeneity, even for single diets commonly used in laboratory studies. More careful consideration of how heterogeneity affects consumer-diet relationships is needed for many field and laboratory systems.     

Beyond carbon and nitrogen: guidelines for estimating three‐dimensional isotopic niche space

Isotopic niche has typically been characterized through carbon and nitrogen ratios and most modeling approaches are limited to two dimensions. Yet, other stable isotopes can provide additional power to resolve questions associated with foraging, migration, dispersal and variations in resource use. The ellipse niche model was recently generalized to n‐dimensions. We present an analogous methodology which incorporates variation across three stable dimensions to estimate the significant features of a population's isotopic niche space including: 1) niche volume (referred to as standard ellipsoid volume, SEV), 2) relative centroid location (CL), 3) shape and 4) area of overlap between multiple ellipsoids and 5) distance between two CLs. We conducted a simulation study showing the accuracy and precision of three dimensional niche models across a range of values. Importantly, the model correctly identifies differences in SEV and CL among populations, even with small sample sizes and in cases where the absolute values cannot precisely be recovered. We use these results to provide guidelines for sample size in conducting multivariate isotopic niche modeling. We demonstrate the utility of our approach with a case study of three bottlenose dolphin populations which appear to possess largely overlapping niches when analyzed with only carbon and nitrogen isotopes. Upon inclusion of sulfur, we see that the three dolphin ecotypes are in fact segregated on the basis of salinity and find the stable isotope niche of inshore bottlenose dolphins significantly larger than coastal and offshore populations.     

蜡质标本制作与保存对沉水植物稳定同位素的影响

实验通过模拟传统方法制作和保存蜡质标本, 旨在探究制作和保存的不同阶段对3种沉水植物碳、氮稳定同位素值的影响。结果表明, 制作和保存标本的过程均会对沉水植物的碳、氮稳定同位素值产生影响。植物标本压制的过程显著降低了沉水植物平均碳稳定同位素值, 其中马来眼子菜下降了1.78‰, 穗花狐尾藻下降了0.98‰, 金鱼藻下降了0.72‰。在压制过程中氮稳定同位素值只有金鱼藻显著下降了1.1‰。沉水植物标本制作的浸泡过程使穗花狐尾藻和马来眼子菜的碳稳定同位素值分别出现了1.00‰和0.52‰的上升, 金鱼藻的碳稳定同位素值增长小于0.08‰, 氮稳定同位素值没有显著变化。沉水植物对标本制作和保存的过程响应存在种间差异。在制作和保存过程中对植物碳稳定同位素产生的影响大于氮稳定同位素值。因此建议, 如果所研究对象的稳定同位素生态学差异小于3‰, 蜡质标本的样品需要进行实验校准, 不然会对结果产生影响。     

Stable Isotope Analysis in Primatology: A Critical Review

Stable isotope analysis has become an important tool in ecology over the last 25 years. A wealth of ecological information is stored in animal tissues in the relative abundances of the stable isotopes of several elements, particularly carbon and nitrogen, because these isotopes navigate through ecological processes in predictable ways. Stable carbon and nitrogen isotopes have been measured in most primate taxonomic groups and have yielded information about dietary content, dietary variability, and habitat use. Stable isotopes have recently proven useful for addressing more fine‐grained questions about niche dynamics and anthropogenic effects on feeding ecology. Here, we discuss stable carbon and nitrogen isotope systematics and critically review the published stable carbon and nitrogen isotope data for modern primates with a focus on the problems and prospects for future stable isotope applications in primatology. Am. J. Primatol. 74:969‐989, 2012. © 2012 Wiley Periodicals, Inc.     

Spatial analysis of stable isotope data to determine primary sources of nutrition for fish

Carbon and nitrogen stable isotopes were used to determine the ultimate autotrophic sources supporting production of three commercially important fish species over unvegetated mudflats in a subtropical estuary. Mean isotope values over the whole estuary for fish and autotroph sources were modeled to indicate feasible combinations of sources. Variability in isotope values among nine locations (separated by 3-10 km) was then used as a further test of the likelihood that sources were involved in fish nutrition. A positive spatial correlation between isotope values of a fish species and an autotroph indicates a substantial contribution from the autotroph. Spatial correlations were tested with a newly developed randomization procedure using differences between fish and autotroph values at each location, based on carbon and nitrogen isotopes combined in two-dimensional space. Both whole estuary modeling and spatial analysis showed that seagrass, epiphytic algae and particulate organic matter in the water column, including phytoplankton, are likely contributors to bream (Acanthopagrus australis) nutrition. However, spatial analysis also showed that mangroves were involved (up to 33% contribution), despite a very low contribution from whole estuary modeling. Spatial analysis on sand whiting (Sillago ciliata) demonstrated the importance of two sources, mangroves (up to 25%) and microalgae on the mudflats, considered unimportant based on whole estuary modeling. No spatial correlations were found between winter whiting (Sillago maculata) and autotrophs, either because fish moved among locations or relied on different autotrophs at different locations. Spatial correlations between consumer and source isotope values provide a useful analytical tool for identifying the role of autotrophs in foodwebs, and demonstrated here that both in situ production of microalgae and organic matter from adjacent habitats were important to fish over mudflats.     

Explaining geographical variation in the isotope composition of mouse lemurs (Microcebus)

Aim We sought to quantify geographical variation in the stable isotope values of mouse lemurs (Microcebus) and to determine whether this variation reflects trophic differences among populations or baseline isotopic differences among habitats. If the latter pattern is demonstrated, then Microcebus can become a proxy for tracking baseline habitat isotopic variability. Establishing such a baseline is crucial for identifying niche partitioning in modern and ancient communities. Location We studied five species of Microcebus from eight distinct habitats across Madagascar. Methods We compared isotopic variation in C₃ plants and Microcebus fur within and among localities. We predicted that carbon and nitrogen isotope values of Microcebus should: (1) vary as a function of abiotic variables such as rainfall and temperature, and (2) covary with isotopic values in plants. We checked for trophic differences among Microcebus populations by comparing the average difference between mouse lemur and plant isotope values for each locality. We then used multiple regression models to explain spatial isotope variation in mouse lemurs, testing a suite of explanatory abiotic variables. Results We found substantial isotopic variation geographically. Ranges for mean isotope values were similar for both Microcebus and plants across localities (carbon 3.5–4.0‰; nitrogen 10.5–11.0‰). Mean mouse lemur and plant isotope values were lowest in cool, moist localities and highest in hot, dry localities. Rainfall explained 58% of the variation in Microcebus carbon isotope values, and mean plant nitrogen isotope values explained 99.7% of the variation in Microcebus nitrogen isotope values. Average differences between mouse lemur and plant isotope values (carbon 5.0‰; nitrogen 5.9‰) were similar across localities. Main conclusions Isotopic data suggest that trophic differences among Microcebus populations were small. Carbon isotope values in mouse lemurs were negatively correlated with rainfall. Nitrogen isotope values in Microcebus and plants covaried. Such findings suggest that nitrogen isotope values for Microcebus are a particularly good proxy for tracking baseline isotopic differences among habitats. Our results will facilitate future comparative research on modern mouse lemur communities, and ecological interpretations of extinct Holocene communities.     

Differential mesowear in occluding upper and lower molars: opening mesowear analysis for lower molars and premolars in hypsodont horses

A new approach of reconstructing ungulate diet, the mesowear method, was recently introduced by Fortelius and Solounias ([2000] Am Mus Novitat 3301:1-36). Mesowear is based on facet development on the occlusal surfaces of the teeth. Restricting mesowear investigation to maxillary cheek teeth would allow mesowear investigation only in assemblages of large numbers of individuals and therefore would generally restrict this method to relatively few assemblages of recent and fossil ungulates. Most of the fossil, subfossil, and recent ungulate osteological assemblages that may be assigned to a single taxon have smaller numbers of individuals. This results in a demand to extend the mesowear method to further tooth positions in order to obtain stable dietary classifications of fossil taxa. The focus of this article is to test if a consistent mesowear classification is obtainable for mandibular as well as for maxillary teeth. For statistical testing, large assemblages of isolated cheek teeth of the Vallesian hipparionine horse Hippotherium primigenium and of the recent zebra Equus burchelli were employed as models. The upper tooth positions P4, M1, M2, and M3 as suggested by Kaiser and Solounias (2003) as the model for the "extended" mesowear method and the lower tooth positions P4-M3 were tested for their consistency in classification of the mesowear variables. We found a considerable shift of the mesowear signature towards the grazing edge of the mesowear continuum in lower cheek teeth. In order to adjust the signal of lower teeth to the signal of the upper teeth, a calibration factor was introduced which allowed incorporation of lower cheek teeth into the same model of mesowear investigation together with upper cheek teeth. We propose that this model is particularly suited for the reconstruction of paleodiets in hypsodont hipparionine and equine equids. We further investigated the functional relation between the mesowear profiles and the distribution of dental tissues along the course of the occlusal contact. We therefore correlated mesowear profiles with enamel distribution profiles and found the mesowear profile to be strongly controlled by the attritional environment encountered by a given apex area. The differential signal observed in cusp apex morphology between upper and lower cheek teeth was found to be more closely related to attrition by the antagonistic tooth than to the distribution of dental tissues in the tooth under consideration. The results suggest a general extension of the mesowear method of paleodiet reconstruction and a basic scenario for the evolution of anisodont dentitions.     

Effects of nutritional restriction on nitrogen and carbon stable isotopes in growing seabirds

When using stable isotopes as dietary tracers it is essential to consider effects of nutritional state on isotopic fractionation. While starvation is known to induce enrichment of ¹⁵N in body tissues, effects of moderate food restriction on isotope signatures have rarely been tested. We conducted two experiments to investigate effects of a 50–55% reduction in food intake on δ¹⁵N and δ¹³C values in blood cells and whole blood of tufted puffin chicks, a species that exhibits a variety of adaptive responses to nutritional deficits. We found that blood from puffin chicks fed ad libitum became enriched in ¹⁵N and ¹³C compared to food-restricted chicks. Our results show that ¹⁵N enrichment is not always associated with food deprivation and argue effects of growth on diet–tissue fractionation of nitrogen stable isotopes (Δ¹⁵N) need to be considered in stable isotope studies. The decrease in δ¹³C of whole blood and blood cells in restricted birds is likely due to incorporation of carbon from ¹³C-depleted lipids into proteins. Effects of nutritional restriction on δ¹⁵N and δ¹³C values were relatively small in both experiments (δ¹⁵N: 0.77 and 0.41‰, δ¹³C: 0.20 and 0.25‰) compared to effects of ecological processes, indicating physiological effects do not preclude the use of carbon and nitrogen stable isotopes in studies of seabird ecology. Nevertheless, our results demonstrate that physiological processes affect nitrogen and carbon stable isotopes in growing birds and we caution isotope ecologists to consider these effects to avoid drawing spurious conclusions.     

Small‐mammal isotope ecology tracks climate and vegetation gradients across western North America

Stable carbon, nitrogen, hydrogen and oxygen isotopes have been used to infer aspects of species ecology and environment in both modern ecosystems and the fossil record. Compared to large mammals, stable isotopic studies of small‐mammal ecology are limited; however, high species and ecological diversity within small mammals presents several advantages for quantifying resource use and organism–environment interactions using stable isotopes over various spatial and temporal scales. We analyzed the isotopic composition of hair from two heteromyid rodent species, Dipodomys ordii and Perognathus parvus, from localities across western North America in order to characterize dietary variation in relation to vegetation and climatic gradients. Significant correlations between the carbon isotopic composition (δ¹³C) of these species and several climatic variables imply that seasonal temperature and precipitation control the composition and distribution of dietary resources (grass seeds). Our results also suggest a moisture influence on the nitrogen isotopic composition (δ¹⁵N) of heteromyid diets. Population‐ and species‐level variation in δ¹³C and δ¹⁵N values record fine‐scale habitat heterogeneity and significant differences in resource use between species. Using classification and regression‐tree techniques, we modeled the geographic variation in heteromyid δ¹³C_diet values based on 10 climatic variables and generated an isotope landscape model (‘isoscape’). The isoscape predictions for δ¹³C_diet differ from expectations based on observed C₄ distributions and instead indicate that D. ordii and P. parvus record seasonally abundant grass resources, with additional model deviations potentially attributed to geographic variation in dietary selection. The oxygen and hydrogen isotopic composition of D. ordii is enriched relative to local meteoric water and suggests that individuals rely on highly evaporated water sources, such as seed moisture. Based on the climatic influences on vegetation and diet documented in this study, the isotopic composition of small mammals has high potential for recording ecological responses to environmental changes over short and long time scales.     

Reconstructing infant weaning histories at Roman period Kellis, Egypt using stable isotope analysis of dentition

Studies of infant feeding and weaning patterns in past populations that rely on a cross-sectional approach must make the assumption that no infant mortality bias exists. Previous investigations of infant weaning patterns at the Dakhleh Oasis, Egypt, relied on cross-sectional isotope data. In this study, we re-examine this weaning pattern, using a simulated longitudinal approach, which does not require any assumptions regarding potential infant mortality biases. This involves examining the dental isotopic signatures of individuals who survived the weaning process. Stable isotope signatures from juveniles and adults (102 individuals, 297 teeth) were examined to reconstruct the weaning history of those that survived the weaning process. Both deciduous and permanent teeth were sampled. Homogenized enamel and dentin samples were isolated from each tooth and analyzed for delta(13)C(ap) and delta(18)O(ap) from the enamel and delta(15)N(coll) and delta(13)C(coll) from dentin collagen. We investigate differences between in utero versus postbirth, preweaning versus postweaning, and juvenile versus adult stable isotope values as reflected in the dentition. A random permutation procedure was used to test for statistically significant differences in stable isotope values between tooth types. Statistically significant differences were observed in all stable isotopes between permanent and deciduous teeth, and between early and later forming permanent teeth in delta(13)C(ap) and delta(15)N(coll) isotopes. These results indicate dietary change between in utero and postbirth, and changes occurring during the weaning period. These results provide a more comprehensive picture of infant weaning practices at Kellis and provide further support that complete weaning occurred by 3 years of age.     

Equations for Lipid Normalization of Carbon Stable Isotope Ratios in Aquatic Bird Eggs

Stable isotope ratios are biogeochemical tracers that can be used to determine the source of nutrients and contaminants in avian eggs. However, the interpretation of stable carbon ratios in lipid-rich eggs is complicated because ¹³C is depleted in lipids. Variation in ¹³C abundance can therefore be obscured by variation in percent lipids. Past attempts to establish an algebraic equation to correct carbon isotope ratios for lipid content in eggs have been unsuccessful, possibly because they relied partly on data from coastal or migratory species that may obtain egg lipids from different habitats than egg protein. We measured carbon, nitrogen and sulphur stable isotope ratios in 175 eggs from eight species of aquatic birds. Carbon, nitrogen and sulphur isotopes were enriched in lipid-extracted egg samples compared with non extracted egg samples. A logarithmic equation using the C∶N ratio and carbon isotope ratio from the non extracted egg tissue calculated 90% of the lipid-extracted carbon isotope ratios within ±0.5‰. Calculating separate equations for eggs laid by species in different habitats (pelagic, offshore and terrestrial-influenced) improved the fit. A logarithmic equation, rather than a linear equation as often used for muscle, was necessary to accurately correct for lipid content because the relatively high lipid content of eggs compared with muscle meant that a linear relationship did not accurately approximate the relationship between percent lipids and the C∶N ratio. Because lipid extraction alters sulphur and nitrogen isotope ratios (and cannot be corrected algebraically), we suggest that isotopic measurement on bulk tissue followed by algebraic lipid normalization of carbon stable isotope ratio is often a good solution for homogenated eggs, at least when it is not possible to complete separate chemical analyses for each isotope.     

Placing butterflies on the map – testing regional geographical resolution of three stable isotopes in Sweden using the monophagus peacock Inachis io

Stable isotope analyses of tissues have been used to help delineate natal regions and routes of migratory animals. The foundations of such studies are isotopic gradients or differences representing geographic regions and habitat used by the organism that are retained in selected tissues. We sampled peacock butterflies Inachis io on a regional level in southern Sweden to study natural variation and the resolving power of the stable isotope method to delineate individuals from known areas on a smaller scale than has typically been used in previous studies. Hydrogen (δD), carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes were obtained from butterflies at seven different locations in an area of 250×250 km over three years (2002–2004). We found sufficient isotopic differences on this regional scale to delineate approximate origins. Of the three isotopes, deuterium showed good discrimination between sites, carbon isotopes showed weaker differentiation, whereas nitrogen isotopes proved unsuitable for small scale studies in this region due to high and unpredictable variation. We found there was enough variation in δD between years to prevent a general application of the technique to resolve sub‐regional variation. Substantial part of this variation was probably caused by seasonal changes in δD of precipitation. These differences produce significant variation in δD between years in animals having short and variable tissue development times, and are difficult to estimate in natural situations. We conclude that stable isotopes are potentially powerful predictors for studies of migratory butterflies in Europe. However, without good knowledge about the sampled individuals’ previous life‐history, a lot of the natural environmental variation in tissue δD cannot be controlled for. In the case of migratory species, this information is difficult to obtain, making the confidence intervals for prediction of natal areas fairly wide and probably only suitable for longer distance migration.     

Differences in nitrogen and carbon stable isotopes between planktonic and benthic microalgae

We compiled published data on the nitrogen and carbon stable isotope ratios of phytoplankton and benthic microalgae from lentic systems and explored the primary factors determining the isotope values among systems. Also, we investigated seasonal changes in nitrogen stable isotope ratios of phytoplankton and benthic microalgae in the strongly acidic lake, Lake Katanuma, which has only two dominant species, Pinnularia acidojaponica as a benthic diatom and Chlamydomonas acidophila, a planktonic green alga. From the published dataset, it may be concluded that δ¹³C of benthic diatoms were more enriched than those of phytoplankton at the same sites, although the nitrogen isotope of phytoplankton and benthic microalgae were similar. This differences in δ¹³C between benthic microalgae and phytoplankton could be explained by the boundary layer effect. On the other hand, nitrogen isotope values of both benthic microalgae and phytoplankton were primarily controlled by the same environmental factor, and boundary layer effects are not the primary factor determining the nitrogen isotope values of microalgae. Also, we showed temporal dynamics in nitrogen isotopes of benthic and planktonic microalgae species in Lake Katanuma, and the trends of nitrogen isotopes are similar between benthic and planktonic microalgae, as concluded from the published dataset.     

The isotopic ecology of East African mammals

Summary The stable carbon and nitrogen isotope ratios of bone collagen have been used to trace diet and habitat selection of the larger mammals of East Africa. 238 individuals of 43 species from montane forests and grasslands in Kenya and Tanzania have been analyzed. The results show that carbon isotopes discriminate between (1) grazers and browsers in savanna grasslands, (2) forest floor and savanna grassland herbivores and (3) forest floor and forest canopy species. Nitrogen isotopes discriminate between (4) carnivores and herbivores, (5) forest and savanna grassland herbivores, and (6) water-dependent and drought-tolerant herbivores. This technique provides a quantitative approach to assessing long-term habitat and diet selection and the role of resource partitioning in animal community structure.Also Archaeology Program     

水分和氮磷水平对小麦碳同位素分辨率和水分利用效率的影响

 依据盆栽试验数据,利用植物稳定性碳同位素分辨率的理论模型,研究了水分和氮磷营养对小麦叶片碳同位素分辨率(Carbon-isotope discrimination △)的影响。结果表明：水分差异引起碳同位素分辨率较大变异,碳同位素分辨率随土壤相对含水量(Soil relative water content)的提高而提高,在土壤相对含水量为60％～70％条件下碳同位素分辨率最高。缺水时磷水平提高,碳同位素分辨率提高。水分利用效率(Water use efficiency)与碳同位素分辨率关系受土壤水分和养分水平的影响。缺水条件下水分利用效率与碳同位素分辨率之间为负相关,充分供水下为正相关;在低氮水平下的关系不明显,施氮150kg·hm-2时相关性显著。