Stable isotopes of carbon and nitrogen in the study of avian and mammalian trophic ecology

Differential fractionation of stable isotopes of carbon during photosynthesis causes C4 plants and C3 plants to have distinct carbon-isotope signatures. In addition, marine C3 plants have stable-isotope ratios of carbon that are intermediate between C4 and terrestrial C3 plants. The direct incorporation of the carbon-isotope ratio (13C/12C) of plants into consumers' tissues makes this ratio useful in studies of animal ecology. The heavy isotope of nitrogen (15N) is preferentially incorporated into the tissues of the consumer from the diet, which results in a systematic enrichment in nitrogen-isotope ratio (15N/14N) with each trophic level. Consequently, stable isotopes of nitrogen have been used primarily to assess position in food chains. The literature pertaining to the use of stable isotopes of carbon and nitrogen in animal trophic ecology was reviewed. Data from 102 studies that reported stable-isotope ratios of carbon and (or) nitrogen of wild birds and (or) mammals were compiled and analyzed relative to diet, latitude, body size, and habitat moisture. These analyses supported the predicted relationships among trophic groups. Carbon-isotope ratios differed among species that relied on C3, C4, and marine food chains. Likewise, nitrogen-isotope ratios were enriched in terrestrial carnivorous mammals relative to terrestrial herbivorous mammals. Also, marine carnivores that ate vertebrates had nitrogen-isotope ratios that were enriched over the ratios of those that ate invertebrates. Data from the literature also indicated that (i) the carbon-isotope ratio of carnivore bone collagen was inversely related to latitude, which was likely the result of an inverse relationship between the proportion of carbon in the food chain that was fixed by C4 plants and latitude; (ii) seabirds and marine mammals from northern oceans had higher nitrogen-isotope ratios than those from southern oceans; (iii) the nitrogen-isotope ratios of terrestrial mammals that used xeric habitats were higher than the ratios of those that used mesic habitats, indicating that water stress can have important effects on the nitrogen-isotope ratio; (iv) there was no relationship between body mass and nitrogen-isotope ratio for either bone collagen or muscle of carnivores; and (v) there was linear covariation between stable-isotope ratios of carbon and nitrogen in marine food chains (but not in terrestrial C3 or C4 food chains), which is likely a product of increases in carbon-isotope ratio with trophic level in marine food chains. Differences in stable-isotope composition among trophic groups were detected despite variation attributable to geographic location, climate, and analytical techniques, indicating that these effects are large and pervasive. Consequently, as knowledge of the distribution of stable isotopes of carbon and nitrogen increases, they will probably become an increasingly important tool in the study of avian and mammalian trophic ecology.