The influence of diet on the distribution of nitrogen isotopes in animals was investigated by analyzing animals grown in the laboratory on diets of constant nitrogen isotopic composition. 
The isotopic composition of the nitrogen in an animal reflects the nitrogen isotopic composition of its diet. The δ^(15)N values of the whole bodies of animals are usually more positive than those of their diets. Different individuals of a species raised on the same diet can have significantly different δ^(15)N values. The variability of the relationship between the δ^(15)N values of animals and their diets is greater for different species raised on the same diet than for the same species raised on different diets. Different tissues of mice are also enriched in ^(15)N relative to the diet, with the difference between the δ^(15)N values of a tissue and the diet depending on both the kind of tissue and the diet involved. The δ^(15)N values of collagen and chitin, biochemical components that are often preserved in fossil animal remains, are also related to the δ^(15)N value of the diet. 
The dependence of the δ^(15)N values of whole animals and their tissues and biochemical components on the δ^(15)N value of diet indicates that the isotopic composition of animal nitrogen can be used to obtain information about an animal's diet if its potential food sources had different δ^(15)N values. The nitrogen isotopic method of dietary analysis probably can be used to estimate the relative use of legumes vs non-legumes or of aquatic vs terrestrial organisms as food sources for extant and fossil animals. However, the method probably will not be applicable in those modern ecosystems in which the use of chemical fertilizers has influenced the distribution of nitrogen isotopes in food sources. 
The isotopic method of dietary analysis was used to reconstruct changes in the diet of the human population that occupied the Tehuacan Valley of Mexico over a 7000 yr span. Variations in the δ^(15)C and δ^(15)N values of bone collagen suggest that C_4 and/or CAM plants (presumably mostly corn) and legumes (presumably mostly beans) were introduced into the diet much earlier than suggested by conventional archaeological analysis.

Influence of Diet On the Distribtion of Nitrogen Isotopes in Animals

As the largest pool of terrestrial organic carbon, soils interact strongly with atmospheric composition, climate, and land cover change. Our capacity to predict and ameliorate the consequences of global change depends in part on a better understanding of the distributions and controls of soil organic carbon (SOC) and how vegetation change may affect SOC distributions with depth. The goals of this paper are (1) to examine the association of SOC content with climate and soil texture at different soil depths; (2) to test the hypothesis that vegetation type, through patterns of allocation, is a dominant control on the vertical distribution of SOC; and (3) to estimate global SOC storage to 3 m, including an analysis of the potential effects of vegetation change on soil carbon storage. We based our analysis on .2700 soil profiles in three global databases supplemented with data for climate, vegetation, and land use. The analysis focused on mineral soil layers. Plant functional types significantly affected the vertical distribution of SOC. The per- centage of SOC in the top 20 cm (relative to the first meter) averaged 33%, 42%, and 50% for shrublands, grasslands, and forests, respectively. In shrublands, the amount of SOC in the second and third meters was 77% of that in the first meter; in forests and grasslands, the totals were 56% and 43%, respectively. Globally, the relative distribution of SOC with depth had a slightly stronger association with vegetation than with climate, but the opposite was true for the absolute amount of SOC. Total SOC content increased with precipitation and clay content and decreased with temperature. The importance of these controls switched with depth, climate dominating in shallow layers and clay content dominating in deeper layers, possibly due to increasing percentages of slowly cycling SOC fractions at depth. To control for the effects of climate on vegetation, we grouped soils within climatic ranges and compared distributions for vegetation types within each range. The percentage of SOC in the top 20 cm relative to the first meter varied from 29% in cold arid shrublands to 57% in cold humid forests and, for a given climate, was always deepest in shrublands, inter- mediate in grasslands, and shallowest in forests ( P , 0.05 in all cases). The effect of vegetation type was more important than the direct effect of precipitation in this analysis. These data suggest that shoot/root allocations combined with vertical root distributions, affect the distribution of SOC with depth. Global SOC storage in the to p3mo fsoil was 2344 Pg C, or 56% more than the 1502 Pg estimated for the first meter (which is similar to the total SOC estimates of 1500-1600 Pg made by other researchers). Global totals for the second and third meters were 491 and 351 Pg C, and the biomes with the most SOC at 1-3 m depth were tropical evergreen forests (158 Pg C) and tropical grasslands/savannas (146 Pg C). Our work suggests that plant functional types, through differences in allocation, help to control SOC distributions with depth in the soil. Our analysis also highlights the potential importance of vegetation change and SOC pools for carbon sequestration strategies.

The vertical distribution of soil organic carbon and its relation to climate and vegetation

To understand the ecology of migratory animals it is important to link geographic regions used by individuals including breeding, wintering, and intermediate stopover sites. Previous conventional approaches used to track animal movements have relied on extrinsic markers and typically the subsequent recovery of individuals. This approach has generally been inappropriate for most small, or non-game animals. The use of intrinsic markers such as fatty acid profiles, molecular DNA analyses, and the measurement of naturally occurring stable isotopes in animal tissues offer alternative approaches. This paper reviews the use of stable isotope analyses (primarily δ13C, δ15N, δ34S, δD, δ87Sr) to trace nutritional origin and migration in animals. This approach relies on the fact that foodweb isotopic signatures are reflected in the tissues of organisms and that such signatures can vary spatially based on a variety of biogeochemical processes. Organisms moving between isotopically distinct foodwebs can carry with them information on the location of previous feeding. Such an approach has been used to track animal use of inshore versus offshore, marine versus freshwater, terrestrial C3 versus marine, terrestrial mesic versus xeric, and C3 versus C4 or Crassulacean acid metabolism foodwebs. More recently, the use of stable hydrogen isotope analyses (δD) to link organisms to broad geographic origin in North America is based on large-scale isotopic contours of growing-season average δD values in precipitation. This technique, especially when combined with the assay of other stable isotopes, will be extremely useful in helping to track migration and movement of a wide range of animals from insects to birds and mammals. Future research to refine our understanding of natural and anthropogenic-induced isotopic gradients in nature, and to explore the use of stable isotopes of other elements, is recommended.

Tracing origins and migration of wildlife using stable isotopes: a review

Differential fractionation of stable isotopes of carbon during photosynthesis causes C 4 plants and C3 plants to have distinct carbon-isotope signatures. In addition, marine C 3 plants have stable-isotope ratios of carbon that are intermediate between C4 and terrestrial C3 plants. The direct incorporation of the carbon-isotope ratio ( 13 C/ 12 C) of plants into consumers' tissues makes this ratio useful in studies of animal ecology. The heavy isotope of nitrogen ( 15 N) is preferentially incorporated into the tissues of the consumer from the diet, which results in a systematic enrichment in nitrogen-isotope ratio ( 15 N/ 14 N) with each trophic level. Consequently, stable isotopes of nitrogen have been used pri- marily 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 ,C 4, and marine food chains. Likewise, nitrogen-isotope ratios were enriched in terres- trial 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 C 4 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 C 3 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. 27 Resume : Le fractionnement differentiel des isotopes stables de carbone durant la photosynthese fait que les plantes C 4 et les plantes C3 ont des signatures d'isotopes de carbone differentes. En outre, chez les plantes C 3 marines, le rapport entre les isotopes stables de carbone est intermediaire entre celui des plantes C 4 et celui des plantes C3 terrestres.

https://www.fs.fed.us/rm/pubs_other/rmrs_2000_kelly_j002.pdf

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

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands (r2 = 0.48) and forests (r2 = 0.17) and for total root biomass in shrublands (r2 = 0.55). On the basis of the best-fit exponential model, the Q10 for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature (r2 = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1469-8137.2000.00681.x

Global patterns of root turnover for terrestrial ecosystems

To determine whether stable isotopes can be used for identifying the geographic origins of migratory bird populations, we examined the isotopic composition of hydrogen (deuterium, δD), carbon (δ13C), and strontium (δ87Sr) in tissues of a migratory passerine, the black-throated blue warbler (Dendroica caerulescens), throughout its breeding range in eastern North America. δD and δ13C values in feathers, which are grown in the breeding area, varied systematically along a latitudinal gradient, being highest in samples from the southern end of the species’ breeding range in Georgia and lowest in southern Canada. In addition, δD decreased from east to west across the northern part of the breeding range, from New Brunswick to Michigan. δ87Sr ratios were highest in the Appalachian Mountains, and decreased towards the west. These patterns are consistent with geographical variation in the isotopic composition of the natural environment, i.e., with that of precipitation, plants, and soils for δD, δ13C, and δ87Sr, respectively. Preliminary analyses of the δD and δ13C composition of feathers collected from warblers in their Caribbean winter grounds indicate that these individuals were mostly from northern breeding populations. Furthermore, variances in isotope ratios in samples from local areas in winter tended to be larger than those in summer, suggesting that individuals from different breeding localities may mix in winter habitats. These isotope markers, therefore, have the potential for locating the breeding origins of migratory species on their winter areas, for quantifying the degree of mixing of breeding populations on migratory and wintering sites, and for documenting other aspects of the population structure migratory animals – information needed for studies of year-round ecology of these species as well as for their conservation. Combining information from several stable isotopes will help to increase the resolution for determining the geographic origins of individuals in such highly vagile populations.

The use of isotope tracers for identifying populations of migratory birds

We used the natural abundance of stable isotopes (carbon and hydrogen) in the feathers of a neotropical migrant songbird to determine where birds from particular breeding areas spend the winter and the extent to which breeding populations mix in winter quarters. We show that most birds wintering on western Caribbean islands come from the northern portion of the species' North American breeding range, whereas those on more easterly islands are primarily from southern breeding areas. Although segregated by breeding latitude, birds within local wintering areas derive from a wide range of breeding longitudes, indicating considerable population mixing with respect to breeding longitude. These results are useful for assessing the effects of wintering habitat loss on breeding population abundances and for predicting whether the demographic consequences will be concentrated or diffuse.

/pdf/linking-breeding-and-wintering-ranges-of-a-migratory-4spx9w9uib.pdf

Linking breeding and wintering ranges of a migratory songbird using stable isotopes

Over the past century, overgrazing and drought in New Mexico’s Jornada Basin has promoted the replacement of native black grama (Bouteloua eriopoda Torr.) grass communities by shrubs, primarily mesquite (Prosopis glandulosa Torr.). We investigated the effects of shrub expansion on the distribution, origin, turnover, and quality of light (LFC) and heavy (HFC) soil organic matter (SOM) fractions using δ13C natural abundance to partition SOM into C4 (grass) and C3 (shrub) sources. Soil organic matter beneath grasses and mesquite was isotopically distinct from associated plant litter, providing evidence of both recent shrub expansion and Holocene plant community changes. Our δ13C analyses indicated that SOM derived from mesquite was greatest beneath shrub canopies, but extended at least 3 m beyond canopy margins, similar to the distribution of fine roots. Specific 14C activities of LFC indicated that root litter is an important source of SOM at depth. Comparison of turnover rates for surface LFC pools in grass (7 or 40 years) and mesquite (11 or 28 years) soils and for HFC pools by soil depth (∼150–280 years), suggest that mesquite may enhance soil C storage relative to grasses. We conclude that the replacement of semiarid grasslands by woody shrubs will effect changes in root biomass, litter production, and SOM cycling that influence nutrient availability and long-term soil C sequestration at the ecosystem level.

Carbon isotopes reveal soil organic matter dynamics following arid land shrub expansion

The cause(s) for the declines in marine mammal populations in the North Pacific and Bering Sea over the past 30 yr are unknown, despite progress in understanding the present ecology of this system. Explanations that attribute the declines to long-term decreases in marine productivity and/or short-term shifts in oceanographic conditions have been offered, but few studies have explored the issue from an explicitly historical perspective. Here we present a high-resolution, 52 yr time series of δ 13 C and δ 15 N values for teeth of juvenile male northern fur seal Callorhinus ursinus collected from a single colony on Saint Paul Island, Alaska in the eastern Bering Sea. Mean annual δ 13 C declined by ~1.1‰ from 1948 to 2000, while long-term mean annual δ 15 N did not significantly change. The relatively small but significant long-term decrease in δ 13 C most likely reflects anthro- pogenically-driven changes in surface ocean carbon reservoirs and not a decline in primary produc- tivity in the North Pacific and Bering Sea system. To assess short-term shifts in the time series, we de- trended the δ 13 C data; the δ 15 N time series did not require a correction. The corrected δ 13 C and uncorrected δ 15 N values showed low amplitude oscillations with a frequency of ~20 to 25 yr that are roughly in phase through time. The relative timing of these oscillations suggest they may be driven by shifts in the Pacific Decadal Oscillation (PDO); however, the frequency of cycles in our time series is approximately half as long as the frequency of the PDO. Finally, mean δ 13 C and δ 15 N values sug- gest that juvenile males modify their migration patterns by Age 3 (GL3, the third year of growth). Specifically, they remain in high-latitude waters year-round (southern Bering Sea and/or western Gulf of Alaska).

/pdf/historic-decline-in-primary-productivity-in-western-gulf-of-4w8jwjzuit.pdf

C. P. Chamberlain

Papers

The use of isotope tracers for identifying populations of migratory birds

Linking breeding and wintering ranges of a migratory songbird using stable isotopes

Carbon isotopes reveal soil organic matter dynamics following arid land shrub expansion

Historic decline in primary productivity in western Gulf of Alaska and eastern Bering Sea: isotopic analysis of northern fur seal teeth