We tested the competing hypotheses that (1) nitrogen discrimination in mammals and birds increases with dietary nitrogen concentration or decreasing C:N ratios and, therefore, discrimination will increase with trophic level as carnivores ingest more protein than herbivores and omnivores or (2) nitrogen discrimination increases as dietary protein quality decreases and, therefore, discrimination will decrease with trophic level as carnivores ingest higher quality protein than do herbivores. Discrimination factors were summarized for five major diet groupings and 21 different species of birds and mammals. Discrimination did not differ between mammals and birds and decreased as protein quality (expressed as biological value) increased with trophic level (i.e., herbivores to carnivores). Relationships between discrimination factors and dietary nitrogen concentration or C:N ratios were either the opposite of what was hypothesized or non-significant. Dietary protein quality accounted for 72% of the variation in discrimination factors across diet groupings. We concluded that protein quality established the baseline for discrimination between dietary groupings, while other variables, such as dietary protein intake relative to animal requirements, created within-group variation. We caution about the care needed in developing studies to understand variation in discrimination and subsequently applying those discrimination factors to estimate assimilated diets of wild animals.

Whitebark pine (Pinus albicaulis) is a masting species that produces relatively large, fat-and protein-rich nuts that are consumed by grizzly bears (Ursus arctos horribilis). Trees produce abundant nut crops in some years and poor crops in other years. Grizzly bear survival in the Greater Yellowstone Ecosystem is strongly linked to variation in pine-nut availability. Because whitebark pine trees are infected with blister rust (Cronartium ribicola), an exotic fungus that has killed the species throughout much of its range in the northern Rocky Mountains, we used stable isotopes to quantify the importance of this food resource to Yellowstone grizzly bears while healthy populations of the trees still exist. Whitebark pine nuts have a sulfur-isotope signature (9.2 ± 1.3‰ (mean ± 1 SD)) that is distinctly different from those of all other grizzly bear foods (ranging from 1.9 ± 1.7‰ for all other plants to 3.1 ± 2.6‰ for ungulates). Feeding trials with captive grizzly bears were used to develop relationships between dietary sulfur-, carbon-, and nitrogenisotope signatures and those of bear plasma. The sulfur and nitrogen relationships were used to estimate the importance of pine nuts to free-ranging grizzly bears from blood and hair samples collected between 1994 and 2001. During years of poor pine-nut availability, 72% of the bears made minimal use of pine nuts. During years of abundant cone availability, 8 ± 10% of the bears made minimal use of pine nuts, while 67 ± 19% derived over 51% of their assimilated sulfur and nitrogen (i.e., protein) from pine nuts. Pine nuts and meat are two critically important food resources for Yellowstone grizzly bears.Résumé : Le pin albicaule (Pinus albicaulis) est un arbre à glandée qui produit des noix relativement grosses et riches en graisses et en protéines, dont se nourrissent les grizzlis (Ursus arctos horribilis). Les arbres donnent des récoltes abondantes certaines années, pauvres d'autres années. La survie des grizzlis de l'écosystème du Grand Yellowstone est étroitement liée aux variations de la disponibilité des noix de pin. Le pin albicaule est sujet aux infections de rouille vésiculaire du pin (Cronartium ribicola), un champignon exotique qui a décimé les populations sur une grande partie de la répartition de l'espèce, dans le nord des Rocheuses; c'est pourquoi nous avons utilisé des isotopes stables pour quantifier l'importance de cette ressource alimentaire pour les grizzlis tandis qu'il existe encore des populations de pins en bonne santé. Les noix du pin albicaule ont une signature d'isotopes du soufre de 9,2 ± 1,3 ‰ (moyenne ± 1 écart type) qui diffère de celles que l'on retrouve dans les autres aliments du grizzli dont les signatures de soufre se situent entre 1,9 ± 1,7 ‰ pour les autres plantes et 3,1 ± 2,6 ‰ pour les ongulés. Des expériences alimentaires sur des grizzlis en captivité ont servi à identifier les relations entre les signatures d'isotopes du soufre, du carbone et de l'azote dans la nourriture et celles du plasma des ours. Les relations du soufre et d...

Accurately predicting isotopic discrimination is central to estimating assimilated diets of wild animals when using stable isotopes. Current mixing models assume that the stable N isotope ratio (delta(15)N) discrimination (Delta(15)N) for each food in a mixed diet is constant and independent of other foods being consumed. Thus, the discrimination value for the mixed diet is the combined, weighted average for each food when consumed as the sole diet. However, if protein quality is a major determinant of Delta(15)N, discrimination values for mixed diets may be higher or lower than the weighted average and will reflect the protein quality of the entire diet and not that of the individual foods. This potential difference occurs because the protein quality of a mixed diet depends on whether, and to what extent, the profiles and amounts of essential amino acids in the individual foods are complementary or non-complementary to each other in meeting the animal's requirement. We tested these ideas by determining the Delta(15)N of several common foods (corn, wheat, alfalfa, soybean, and fish meal) with known amino acid profiles when fed singly and in combination to laboratory rats. Discrimination values for the mixed diets often differed from the weighted averages for the individual foods and depended on the degree of complementation. Delta(15)N for mixed diets ranged from 1.1 per thousand lower than the weighted average for foods with complementary amino acid profiles to 0.4 per thousand higher for foods with non-complementary amino acid profiles. These differences led to underestimates as high as 44% and overestimates as high as 36% of the relative proportions of fish meal and soybean meal N, respectively, in the assimilated mixed diets. We conclude that using isotopes to estimate assimilated diets is more complex than often appreciated and will require developing more biologically based, time-sensitive models.

Summary1. Accurately predicting isotope ratio discrimination is central to using mixing models to estimate assimilated diets of wild animals. This process is complicated when omnivores consume mixed diets because their discrimination is unlikely to be the weighted average of the various dietary constituents as occurs in current models. in all diets, sulphur should be highly conserved as indicated by the lack of any change in D 34 S when diets were supplemented with carbohydrates and fat to both provide additional energy and reduce protein content. 6. Predictive equations incorporating both protein quality and quantity accounted for 81% of the variation in D 15 N and offer the opportunity to create more realistic mixing models to accurately estimate assimilated diets for omnivores.

Energy maximization, time minimization, and linear programming models subject to various constraints have dominated foraging ecology ideas and methods for decades. However, animals must use very complex physiological processes and foraging decisions to ensure fitness that in many cases may not be adequately described by these approaches. An example of this problem occurs when brown bears, Ursus arctos, have access to both abundant salmon and fruit. Salmon are one of the most energy and nutrient dense foods available to bears. Fruits are often high in soluble carbohydrates, low to deficient in many required nutrients, and more difficult to efficiently exploit. Therefore, wild brown bears that fatten primarily on fruits without access to salmon are 50% smaller than salmon-feeding bears. Thus, we predicted based on a linear, energy-maximizing model without dietary interaction effects that wild brown bears with access to both abundant salmon and fruit would feed almost exclusively on salmon. However, wild adult females with or without accompanying offspring foraged three times longer per day on fruit than on salmon. Similarly, the relative dry matter intake of ad libitum apples and salmon by captive, adult brown bears averaged 7695% fruit and 2495% salmon. Captive brown bears consuming mixed diets with intermediate dietary protein levels had 60% lower maintenance energy costs, 37% to 139% higher efficiencies of mass gain, and 72% to 520% higher maximum rates of gain than when they consumed either salmon or fruit alone. These relationships were nonlinear functions of dietary protein content in which salmon and fruit provided complementary nutritional resources. Both wild and captive bears attempted to regulate total protein, energy, and carbohydrate intake within a multidimensional intake target that both maximized energy intake and mass gain.

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Spawning cutthroat trout (Oncorhynchus clarki (Richardson, 1836)) are a potentially important food resource for grizzly bears (Ursus arctos horribilis Ord, 1815) in the Greater Yellowstone Ecosystem. We developed a method to estimate the amount of cutthroat trout ingested by grizzly bears living in the Yellowstone Lake area. The method utilized (i) the relatively high, naturally occurring concentration of mercury in Yellowstone Lake cutthroat trout (508 ± 93 ppb) and its virtual absence in all other bear foods ([Formula: see text]6 ppb), (ii) hair snares to remotely collect hair from bears visiting spawning cutthroat trout streams between 1997 and 2000, (iii) DNA analyses to identify the individual and sex of grizzly bears leaving a hair sample, (iv) feeding trials with captive bears to develop relationships between fish and mercury intake and hair mercury concentrations, and (v) mercury analyses of hair collected from wild bears to estimate the amount of trout consumed by each bear. Male grizzly bears consumed an average of 5 times more trout/kg bear than did female grizzly bears. Estimated cutthroat trout intake per year by the grizzly bear population was only a small fraction of that estimated by previous investigators, and males consumed 92% of all trout ingested by grizzly bears.