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

Background 
Stable isotope analysis is increasingly being utilised across broad areas of ecology and biology. Key to much of this work is the use of mixing models to estimate the proportion of sources contributing to a mixture such as in diet estimation.


Methodology 
By accurately reflecting natural variation and uncertainty to generate robust probability estimates of source proportions, the application of Bayesian methods to stable isotope mixing models promises to enable researchers to address an array of new questions, and approach current questions with greater insight and honesty.


Conclusions 
We outline a framework that builds on recently published Bayesian isotopic mixing models and present a new open source R package, SIAR. The formulation in R will allow for continued and rapid development of this core model into an all-encompassing single analysis suite for stable isotope research.

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Source partitioning using stable isotopes: coping with too much variation.

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

Measurements of δ15N 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 δ15N enrichment within this general pattern. We conducted an extensive literature review, which yielded 134 estimates from controlled studies of consumer-diet δ15N 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 δ15N 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. δ15N 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 δ15N enrichment. The lower δ15N 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 δ15N 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 δ15N 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 δ15N 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.

Sources of variation in consumer-diet δ15N enrichment: a meta-analysis

Food limitation is an important issue in ecology because it can influence life history traits, population sizes, and community structure (through effects of competition). Work at the level of populations and communities has led to arguments that food limitation and competition are more important in winter than during the breeding season (e.g. 1, 2, 7, 8, 83, 85, 211, 234). In fact, it is commonly argued that food is superabundant during the breeding season (7, 8, 161, 213, 246, 250, 251, 308-311). However, such arguments are based on indirect rather than direct evidence of the effects of food on reproduction and survival (fitness) (149). Direct evidence for food limitation and competition in winter exists when survival is affected, and some experimental evidence suggests such effects (e.g. 75, 113, 122, 132). However, if current or future reproductive success is limited by food, then food limitation will also exist during the breeding season. Reproductive ecologists historically have argued that food limits reproductive success (e.g. 13, 297). Yet, even this school has included recent arguments against food limitation. For instance, Ettinger & King (77) think that perching time of birds commonly reflects loafing time because birds set their clutch and brood sizes based on years and periods of stringent (low food) conditions. However, perching time may not reflect loafing but rather an important time commitment to reproductive success (T. Martin, unpublished ms.) Thus, the status of food limitation in breeding birds is not clear. Here, I review evidence for food limitation in the context of life history theory because it provides a fundamental framework from which to interpret

Food as a limit on breeding birds: a life-history perspective

Studies using measurements of61'5N to delineate diet or trophic level in natural ecosystems are based on the premise that 6'5N values in consumer tissues can be reliably correlated with those in the diet. However, juvenile Japanese Quail (Coturnixjaponica) fed a rationed diet designed to maintain, but not increase, body mass showed significantly enriched tissue 6'5N values over a control group fed the same diet ad libitum. We tested the hypothesis that fasting or nutritional stress can also cause elevated 6'5N values in tissues of wild birds by examining tissues of Arctic-nesting female Ross' Geese (Chen rossii) before and after their period of fasting during egg laying and incubation. Significant declines in body, pectoral muscle, liver and abdominal fat mass occurred from arrival through incubation. Post-incubating geese showed significantly higher pectoral muscle and liver 6'5N values compared to geese taken before clutch initiation but 6a3C values in these tissues were unchanged. We hypothesize a mechanism of tissue 6'5N enrichment due to reduced nutrient intake and discuss the implications of these results to ecosystem studies using stable-nitrogen isotope analysis.

Stable-Nitrogen Isotope Enrichment in Avian Tissues Due to Fasting and Nutritional Stress: Implications for Isotopic Analyses of Diet

The Mid-Continent Population of the lesser snow goose, which breeds in the eastern and central Canadian Arctic and sub-Arctic, and winters in the southern United States and northern Mexico has increased 5–7% annually from the late 1960s to the mid-1990s, largely because of increased survival in response to an agricultural food subsidy. The rise in numbers complements the increased use of nitrogen fertilizers and a corresponding rise in yields of rice, corn, and wheat along the flyways and on the wintering grounds. In sub-Arctic migration areas and at Arctic breeding colonies, foraging by high numbers of birds has led to loss of coastal vegetation, adverse changes in soil properties and the establishment of an alternative stable state of exposed sediment, which can be detected with LANDSAT imagery. At a local scale, gosling growth, size and survival decreased in affected areas and other taxa have been adversely affected. The food subsidy on wintering and migration areas appears insufficient to meet reproductive demands as foraging in spring continues to occur on southern Hudson Bay staging and nesting areas. The recent introduction of liberal hunting regulations may reduce population size in the near term, but the revegetation of these coastal ecosystems will take decades to achieve. The present pattern of vegetation loss in these Arctic coastal systems is likely to continue in the forseeable future.

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The dynamics of landscape change and snow geese in mid‐continent North America

We employ mitochondrial (mt) DNA markers to examine the matrilineal component of population genetic structure in the snow goose Chen caerulescens. From banding returns, it is known that females typically nest at their natal or prior nest site, whereas males pair with females on mixed wintering grounds and mediate considerable nuclear gene flow between geographically separate breeding colonies. Despite site philopatry documented for females, mtDNA markers show no clear distinctions between nesting populations across the species' range from Wrangel Island, USSR to Baffin Island in the eastern Canadian Arctic. Two major mtDNA clades (as well as rare haplotypes) are distributed widely and provide one of the few available examples of a phylogeographic pattern in which phylogenetic discontinuity in a gene tree exists without obvious geographic localization within a species' range. The major mtDNA clades may have differentiated in Pleistocene refugia, and colonized current nesting sites through recent range expansion via pulsed or continual low-level dispersal by females. The contrast between results of banding returns and mtDNA distributions in the snow goose raises general issues regarding population structure: direct contemporary observations on dispersal and gene flow can in some cases convey a misleading impression of phylogeographic population structure, because they fail to access the evolutionary component of population connectedness; conversely, geographic distributions of genetic markers can provide a misleading impression of contemporary dispersal and gene flow because they retain a record of evolutionary events and past demographic parameters that may differ from those of the present. An understanding of population structure requires integration of both evolutionary (genetic) and contemporary (direct observational) perspectives.

/pdf/matriarchal-population-genetic-structure-in-an-avian-species-1fsslf0rhj.pdf

Matriarchal population genetic structure in an avian species with female natal philopatry.

We assessed the effectiveness of an extensive and unprecedented wildlife reduction effort directed at a wide-ranging migratory population of geese. Population reduction efforts that targeted several populations of light geese (greater snow geese (Chen caerulescens atlantica), lesser snow geese (C. c. caerulescens), and Ross's geese (C. rossii)) began in 1999 in central and eastern North America. Such efforts were motivated by a broad consensus that abundance of these geese was causing serious ecological damage to terrestrial and salt marsh ecosystems in central and eastern parts of the Canadian Arctic and subarctic regions along Hudson Bay. Starting in February 1999, special conservation measures (or, in the U.S., a conservation order) were added to the respective federal regulations that permitted hunters to take snow geese (in parts of Canada and the U.S.) and Ross's geese (in parts of the U.S.) during specified harvest periods outside of the hunting season. These measures were accompanied by increase or removal of daily kill and possession limits and by permissions to use previously prohibited equipment for hunting these species in certain regions of the continent. The intent was to reduce adult survival through increased hunting mortality, which was judged to be the most cost-effective approach to reversing population growth. Our principal goal was to assess the effectiveness of reduction efforts directed at the midcontinent population of lesser snow geese, which was thought to be the most serious threat to arctic and subarctic ecosystems of the 3 light goose populations. Our multiple objectives included the estimation and detection of change in the response measures of total annual harvest, harvest rate, survival rate, and abundance, using the 1998 hunting period (defined as 1 Aug 1998 to 31 Jul 1999) as a point of reference. We used information about hunter recoveries of leg- banded snow geese and estimates of regular-season harvest to estimate 1) conservation-order harvest and total annual harvest, 2) geographic and temporal distribution of recoveries by age class, 3) survival and recovery probability, and 4) abundance of snow geese each August using Lincoln's (1930) method. We also modeled population growth to infer the form of population response to management efforts. Toward that end, we also proposed a method of estimating conservation-order harvest and tested for differences in band-reporting rate between Canada and the United States. Overall, the balance of evidence favored the conclusion that the midcontinent population has continued to grow during the conservation order, although perhaps at a reduced rate. We suggest that annual rate of population growth ð^ lÞ, derived from estimates of annual population size in August, likely provides the most reliable inference about change in the midcontinent population. There was a decline in annual survival probability between these 2 periods from about 0.89 to about 0.83 among snow geese from the southern-nesting stratum (south of 608N latitude), thought to compose about 10% of the midcontinent population. However, we detected no change in the much larger northern-nesting stratum (north of 608N latitude), where annual survival remained at about 0.87 from 1989 to 2006. Thus, the conclusion that this population continued to increase during the conservation order was largely consistent with the finding that a weighted-survival probability for midcontinent snow geese essentially did not change between the period preceding (1989-1997) and during (1998-2006) the conservation order. Consistent with high survival rates were low harvest rates, which increased from 0.024 during 1989-1997 for northern geese to only 0.027 during 1998-2006 and from 0.031 to only 0.037 for southern geese. Despite the initial increase associated with the conservation order, harvest rates declined during the conservation order for geese from both strata. We suggest that the higher harvest rate evident for

/pdf/harvest-survival-and-abundance-of-midcontinent-lesser-snow-38et41ocgw.pdf

Ray T. Alisauskas

Papers

Stable-Nitrogen Isotope Enrichment in Avian Tissues Due to Fasting and Nutritional Stress: Implications for Isotopic Analyses of Diet

The dynamics of landscape change and snow geese in mid‐continent North America

Matriarchal population genetic structure in an avian species with female natal philopatry.

Harvest, survival, and abundance of midcontinent lesser snow geese relative to population reduction efforts

The role of nutrient reserves in limiting waterfowl reproduction