Measurements of delta15N of consumers are usually higher than those of their diet. This general pattern is widely used to make inferences about trophic relationships in ecological studies, although the underlying mechanisms causing the pattern are poorly understood. However, there can be substantial variation in consumer-diet delta15N enrichment within this general pattern. We conducted an extensive literature review, which yielded 134 estimates from controlled studies of consumer-diet delta15N enrichment, to test the significance of several potential sources of variation by means of meta-analyses. We found patterns related to processes of nitrogen assimilation and excretion. There was a significant effect of the main biochemical form of nitrogenous waste: ammonotelic organisms show lower delta15N enrichment than ureotelic or uricotelic organisms. There were no significant differences between animals feeding on plant food, animal food, or manufactured mixtures, but detritivores yielded significantly lower estimates of enrichment. delta15N enrichment was found to increase significantly with the C:N ratio of the diet, suggesting that a nitrogen-poor diet can have an effect similar to that already documented for fasting organisms. There were also differences among taxonomic classes: molluscs and crustaceans generally yielded lower delta15N enrichment. The lower delta 15N enrichment might be related to the fact that molluscs and crustaceans excrete mainly ammonia, or to the fact that many were detritivores. Organisms inhabiting marine environments yielded significantly lower estimates of delta15N enrichment than organisms inhabiting terrestrial or freshwater environments, a pattern that was influenced by the number of marine, ammonotelic, crustaceans and molluscs. Overall, our analyses point to several important sources of variation in delta15N enrichment and suggest that the most important of them are the main biochemical form of nitrogen excretion and nutritional status. The variance of estimates of delta15N enrichment, as well as the fact that enrichment may be different in certain groups of organisms should be taken into account in statistical approaches for studying diet and trophic relationships.
Animals reflect the stable isotope ratios (␦ 15 N and ␦ 13 C) of their diet, with a slight enrichment in 15 N that allows the use of ␦ 15 N as a trophic-level indicator. Stable isotope contents were measured for the litter, soil, and macro-invertebrates of three temperate deciduous forest sites, in spring, summer, and autumn, to study the trophic structure of this community. No distinct trophic structure could be derived from measurements of ␦ 13 C. In contrast, when the ␦ 15 N values of all animal species were grouped together, the hypothesis of an isotopically similar diet was rejected. Therefore, the community spreads over more than one trophic level and was subdivided into detritivores and predators. The potential detritivore food sources in the forest litter and soil showed a variety of isotopic ratios. Despite this fact, the variance of the isotopic ratios of the detritivorous species was not larger than what could be expected from interspecific variability of trophic isotopic enrichment alone. This was also the case for the predators in most of the sample sets. However, in some cases this variance was significantly larger, due to a small number of species with high ␦ 15 N values. The ␦ 15 N values of the detritivores indicated that the mean ␦ 15 N value of their food was close to that measured for the superficial litter layers. The difference in ␦ 15 N between detritivores and predators was highly significant and never significantly different from the value expected for one trophic transfer (3.4‰), although often slightly higher. Most of the litter macro-invertebrate community we studied can therefore be described as belonging to two trophic levels, one feeding on the superficial litter layers (or on soil fractions that have a similar ␦ 15 N value), and a second trophic level feeding on the first, with some indication of intraguild predation among the predators.Between-site differences of up to 7‰ were found for ␦ 15 N in the litter, and the ␦ 15 N values of the whole animal community were shifted in accordance with the local value of the litter ␦ 15 N. Therefore, the trophic structure must be studied in relation to the local isotopic content of the litter. Seasonal differences in isotopic ratios of the litter or animal samples were neither large nor consistent. These findings indicate similar trophic structure of the communities at the three sites and during the three sampling periods.
Although a growing body of work supports the plausibility of sympatric speciation in animals, the practical difficulties of directly quantifying reproductive isolation between diverging taxa remain an obstacle to analyzing this process. We used a combination of genetic and biogeochemical markers to produce a direct field estimate of assortative mating in phytophagous insect populations. We show that individuals of the same insect species, the European corn borer Ostrinia nubilalis, that develop on different host plants can display almost absolute reproductive isolation-the proportion of assortative mating was >95%-even in the absence of temporal or spatial isolation.
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