Diet energy density estimated from isotopes in predator hair associated with survival, habitat, and population dynamics.

Sea ice loss is fundamentally altering the Arctic marine environment. Yet there is a paucity of data on the adaptability of food webs to ecosystem change, including predator-prey interactions. Polar bears (Ursus maritimus) are an important subsistence resource for Indigenous people and an apex predator that relies entirely on the under-ice food web to meet its energy needs. In this study, we assessed whether polar bears maintained dietary energy density by prey switching in response to spatiotemporal variation in prey availability. We compared the macronutrient composition of diets inferred from stable carbon and nitrogen isotopes in polar bear guard hair (primarily representing summer/fall diet) during periods when bears had low and high survival (2004-2016), between bears that summered on land versus pack ice, and between bears occupying different regions of the Alaskan and Canadian Beaufort Sea. Polar bears consumed diets with lower energy density during periods of low survival, suggesting that concurrent increased dietary proportions of beluga whales (Delphinapterus leucas) did not offset reduced proportions of ringed seals (Pusa hispida). Diets with the lowest energy density and proportions from ringed seal blubber were consumed by bears in the western Beaufort Sea (Alaska) during a period when polar bear abundance declined. Intake required to meet energy requirements of an average free-ranging adult female polar bear was 2.1 kg/day on diets consumed during years with high survival but rose to 3.0 kg/day when survival was low. Although bears that summered onshore in the Alaskan Beaufort Sea had higher-fat diets than bears that summered on the pack ice, access to the remains of subsistence-harvested bowhead whales (Balaena mysticetus) contributed little to improving diet energy density. Because most bears in this region remain with the sea ice year round, prey switching and consumption of whale carcasses onshore appear insufficient to augment diets when availability of their primary prey, ringed seals, is reduced. Our results show that a strong predator-prey relationship between polar bears and ringed seals continues in the Beaufort Sea. The method of estimating dietary blubber using predator hair, demonstrated here, provides a new metric to monitor predator-prey relationships that affect individual health and population demographics.

Animal tracing with sulfur isotopes: Spatial segregation and climate variability in Africa likely contribute to population trends of a migratory songbird.

Climatic conditions affect animals but range-wide impacts at the population level remain largely unknown, especially in migratory species. However, studying climate-population relationships is still challenging in small migrants due to a lack of efficient and cost-effective geographic tracking method. Spatial distribution patterns of environmental stable isotopes (so called 'isoscapes') generally overcome these limitations but none of the currently available isoscapes provide a substantial longitudinal gradient in species-rich sub-Saharan Africa. In this region, sulphur (δ34 S) has not been sufficiently explored on a larger scale. We developed a δ34 S isoscape to trace animal origins in sub-Saharan Africa by coupling known-origin samples from tracked migratory birds with continental remotely sensed environmental data building on environment-δ34 S relationships using a flexible machine learning technique. Furthermore, we link population-specific nonbreeding grounds with interannual climatic variation that might translate to breeding population trends. The predicted δ34 S isotopic map featured east-west and coast-to-inland isotopic gradients and was applied to predict nonbreeding grounds of three breeding populations of Eurasian Reed Warblers Acrocephalus scirpaceus with two distinct migratory phenotypes. Breeding populations as well as migratory phenotypes exhibited large-scale segregation within the African nonbreeding range. These regions also differed substantially in the interannual climatic variation, with higher interannual variability in the eastern part of the range during 2001-2012. Over the same period, the eastern European breeding population seemed to have experienced a more steep decline in population size. The link between migratory patterns and large-scale climatic variability appears important to better understand population trajectories in many declining migratory animals. We believe animal tracing using sulphur isotopes will facilitate these efforts and offers manifold ecological and forensic applications in the biodiversity hotspot of sub-Saharan Africa.

Complex Life Histories Alter Patterns of Mercury Exposure and Accumulation in a Pond-Breeding Amphibian.

Quantifying how contaminants change across life cycles of species that undergo metamorphosis is critical to assessing organismal risk, particularly for consumers. Pond-breeding amphibians can dominate aquatic animal biomass as larvae and are terrestrial prey as juveniles and adults. Thus, amphibians can be vectors of mercury exposure in both aquatic and terrestrial food webs. However, it is still unclear how mercury concentrations are affected by exogenous (e.g., habitat or diet) vs endogenous factors (e.g., catabolism during hibernation) as amphibians undergo large diet shifts and periods of fasting during ontogeny. We measured total mercury (THg), methylmercury (MeHg), and isotopic compositions (δ 13C, δ15N) in boreal chorus frogs (Pseudacris maculata) across five life stages in two Colorado (USA) metapopulations. We found large differences in concentrations and percent MeHg (of THg) among life stages. Frog MeHg concentrations peaked during metamorphosis and hibernation coinciding with the most energetically demanding life cycle stages. Indeed, life history transitions involving periods of fasting coupled with high metabolic demands led to large increases in mercury concentrations. The endogenous processes of metamorphosis and hibernation resulted in MeHg bioamplification, thus decoupling it from the light isotopic proxies of diet and trophic position. These step changes are not often considered in conventional expectations of how MeHg concentrations within organisms are assessed.

Temporal Influences on Selenium Partitioning, Trophic Transfer, and Exposure in a Major U.S. River.

Hydrologic and irrigation regimes mediate the timing of selenium (Se) mobilization to rivers, but the extent to which patterns in Se uptake and trophic transfer through recipient food webs reflect the temporal variation in Se delivery is unknown. We investigated Se mobilization, partitioning, and trophic transfer along approximately 60 river miles of the selenium-impaired segment of the Lower Gunnison River (Colorado, USA) during six sampling trips between June 2015 and October 2016. We found temporal patterns in Se partitioning and trophic transfer to be independent of those in dissolved Se concentrations and that the recipient food web sustained elevated Se concentrations from earlier periods of high Se mobilization. Using an ecosystem-scale Se accumulation model tailored to the Lower Gunnison River, we predicted that the endangered Razorback Sucker (Xyrauchen texanus) and Colorado Pikeminnow (Ptychocheilus lucius) achieve whole-body Se concentrations exceeding aquatic life protection criteria during periods of high runoff and irrigation activity (April-August) that coincide with susceptible phases of reproduction and early-life development. The results of this study challenge assumptions about Se trophodynamics in fast-flowing waters and introduce important considerations for the management of Se risks for biota in river ecosystems.

Migratory divides coincide with reproductive barriers across replicated avian hybrid zones above the Tibetan Plateau.

Migratory divides are proposed to be catalysts for speciation across a diversity of taxa. However, it is difficult to test the relative contributions of migratory behaviour vs. other divergent traits to reproductive isolation. Comparing hybrid zones with and without migratory divides offers a rare opportunity to directly examine the contribution of divergent migratory behaviour to reproductive barriers. We show that across replicate sampling transects of two pairs of barn swallow (Hirundo rustica) subspecies, strong reproductive isolation coincided with a migratory divide spanning 20 degrees of latitude. A third subspecies pair exhibited no evidence for a migratory divide and hybridised extensively. Within migratory divides, overwintering habitats were associated with assortative mating, implicating a central contribution of divergent migratory behaviour to reproductive barriers. The remarkable geographic coincidence between migratory divides and genetic breaks supports a long-standing hypothesis that the Tibetan Plateau is a substantial barrier contributing to the diversity of Siberian avifauna.

Benthic algal (periphyton) growth rates in response to nitrogen and phosphorus: Parameter estimation for water quality models.

Nitrogen (N) and phosphorus (P) are significant pollutants that can stimulate nuisance blooms of algae. Water quality models (e.g., WASP, CE-QUAL-R1, CE-QUAL-ICM, QUAL2k) are valuable and widely used management tools for algal accrual due to excess nutrients in the presence of other limiting factors. These models utilize the Monod and Droop equations to associate algal growth rate with dissolved nutrient concentration and intra-cellular nutrient content. Having accurate parameter values is essential to model performance, however published values for model parameterization are limited, particularly for benthic (periphyton) algae. We conducted a 10-day mesocosm experiment and measured diatom-dominated periphyton biomass accrual through time as chlorophyll a (chl a) and ash-free dry mass (AFDM) in response to additions of N (range 5-11,995 µg NO3-N/L) and P (range 0.89-59.51 µg SRP/L). Resulting half saturation coefficients and growth rates are similar to other published values, but minimum nutrient quotas are higher than those previously reported. Saturation concentration for N ranged from 150 to 2450 µg NO3-N/L based on chl a and from 8.5 to 60 µg NO3-N/L when based on AFDM. Similarly, the saturation concentration for P ranged from 12 to 29 µg-P/L based on chl a, and from 2.5 to 6.1 µg-P/L based on AFDM. These saturation concentrations provide an upper limit for streams where diatom growth can be expected to respond to nutrient levels and a benchmark for reducing nutrient concentrations to a point where benthic algal growth will be limited.

Metamorphosis Affects Metal Concentrations and Isotopic Signatures in a Mayfly (Baetis tricaudatus): Implications for the Aquatic-Terrestrial Transfer of Metals.

Insect metamorphosis often results in substantial chemical changes that can alter contaminant concentrations and fractionate isotopes. We exposed larval mayflies (Baetis tricaudatus) and their food (periphyton) to an aqueous zinc gradient (3-340 µg Zn/l) and measured zinc concentrations at different stages of metamorphosis: larval, subimago, and imago. We also measured changes in stable isotopes (δ15N and δ13C) in unexposed mayflies. Larval zinc concentrations were positively related to aqueous zinc, increasing 9-fold across the exposure gradient. Adult zinc concentrations were also positively related to aqueous zinc, but were 7-fold lower than larvae. This relationship varied according to adult substage and sex. Tissue concentrations in female imagoes were not related to exposure concentrations, but the converse was true for all other stage-by-sex combinations. Metamorphosis also increased δ15N by ∼0.8‰, but not δ13C. Thus, the main effects of metamorphosis on insect chemistry were large declines in zinc concentrations coupled with increased δ15N signatures. For zinc, this change was largely consistent across the aqueous exposure gradient. However, differences among sexes and stages suggest that caution is warranted when using nitrogen isotopes or metal concentrations measured in one insect stage (e.g., larvae) to assess risk to wildlife that feed on subsequent life stages (e.g., adults).

Reproductive Allochrony in Seasonally Sympatric Populations Maintained by Differential Response to Photoperiod: Implications for Population Divergence and Response to Climate Change.

Reproductive allochrony presents a potential barrier to gene flow and is common in seasonally sympatric migratory and sedentary birds. Mechanisms mediating reproductive allochrony can influence population divergence and the capacity of populations to respond to environmental change. We asked whether reproductive allochrony in seasonally sympatric birds results from a difference in response to supplementary or photoperiodic cues and whether the response varies in relation to the distance separating breeding and wintering locations as measured by stable isotopes. We held seasonally sympatric migratory and sedentary male dark-eyed juncos (Junco hyemalis) in a common garden in early spring under simulated natural changes in photoperiod and made measurements of reproductive and migratory physiology. On the same dates and photoperiods, sedentary juncos had higher testosterone (initial and gonadotropin-releasing hormone induced), more developed cloacal protuberances, and larger testes than migrants. In contrast, migratory juncos had larger fat reserves (fuel for migration). We found a negative relationship between testis mass and feather hydrogen isotope ratios, indicating that testis growth was more delayed in migrants making longer migrations. We conclude that reproductive allochrony in seasonally sympatric migratory and sedentary birds can result from a differential response to photoperiodic cues in a common garden, and as a result, gene flow between migrants and residents may be reduced by photoperiodic control of reproductive development. Further, earlier breeding in response to future climate change may currently be constrained by differential response to photoperiodic cues.

Holocene dynamics of the Florida Everglades with respect to climate, dustfall, and tropical storms.

Aeolian dust is rarely considered an important source for nutrients in large peatlands, which generally develop in moist regions far from the major centers of dust production. As a result, past studies assumed that the Everglades provides a classic example of an originally oligotrophic, P-limited wetland that was subsequently degraded by anthropogenic activities. However, a multiproxy sedimentary record indicates that changes in atmospheric circulation patterns produced an abrupt shift in the hydrology and dust deposition in the Everglades over the past 4,600 y. A wet climatic period with high loadings of aeolian dust prevailed before 2800 cal BP (calibrated years before present) when vegetation typical of a deep slough dominated the principal drainage outlet of the Everglades. This dust was apparently transported from distant source areas, such as the Sahara Desert, by tropical storms according to its elemental chemistry and mineralogy. A drier climatic regime with a steep decline in dustfall persisted after 2800 cal BP maintaining sawgrass vegetation at the coring site as tree islands developed nearby (and pine forests covered adjacent uplands). The marked decline in dustfall was related to corresponding declines in sedimentary phosphorus, organic nitrogen, and organic carbon, suggesting that a close relationship existed between dustfall, primary production, and possibly, vegetation patterning before the 20th century. The climatic change after 2800 cal BP was probably produced by a shift in the Bermuda High to the southeast, shunting tropical storms to the south of Florida into the Gulf of Mexico.

Individual specialization in the foraging habits of female bottlenose dolphins living in a trophically diverse and habitat rich estuary.

We examine individual specialization in foraging habits (foraging habitat and trophic level) of female bottlenose dolphins (Tursiops truncatus) resident in Sarasota Bay, Florida, USA, by analyzing time series of stable isotope (δ(15)N and δ(13)C) values in sequential growth layer groups within teeth. The isotope data provide a chronology of foraging habits over the lifetime of the individual and allowed us to show that female bottlenose dolphins exhibit a high degree of individual specialization in both foraging habitat and trophic level. The foraging habits used by adult females are similar to those they used as calves and may be passed down from mother to calf through social learning. We also characterized the foraging habits and home range of each individual by constructing standard ellipses from isotope values and dolphin sightings data (latitude and longitude), respectively. These data show that Sarasota Bay bottlenose dolphins forage within a subset of the habitats in which they are observed. Moreover, females with similar observational standard ellipses often possessed different foraging specializations. Female bottlenose dolphins may demonstrate individual specialization in foraging habits because it reduces some of the cost of living in groups, such as competition for prey.

Continental-scale, seasonal movements of a heterothermic migratory tree bat.

Long-distance migration evolved independently in bats and unique migration behaviors are likely, but because of their cryptic lifestyles, many details remain unknown. North American hoary bats (Lasiurus cinereus cinereus) roost in trees year-round and probably migrate farther than any other bats, yet we still lack basic information about their migration patterns and wintering locations or strategies. This information is needed to better understand unprecedented fatality of hoary bats at wind turbines during autumn migration and to determine whether the species could be susceptible to an emerging disease affecting hibernating bats. Our aim was to infer probable seasonal movements of individual hoary bats to better understand their migration and seasonal distribution in North America. We analyzed the stable isotope values of non-exchangeable hydrogen in the keratin of bat hair and combined isotopic results with prior distributional information to derive relative probability density surfaces for the geographic origins of individuals. We then mapped probable directions and distances of seasonal movement. Results indicate that hoary bats summer across broad areas. In addition to assumed latitudinal migration, we uncovered evidence of longitudinal movement by hoary bats from inland summering grounds to coastal regions during autumn and winter. Coastal regions with nonfreezing temperatures may be important wintering areas for hoary bats. Hoary bats migrating through any particular area, such as a wind turbine facility in autumn, are likely to have originated from a broad expanse of summering grounds from which they have traveled in no recognizable order. Better characterizing migration patterns and wintering behaviors of hoary bats sheds light on the evolution of migration and provides context for conserving these migrants.

Metamorphosis alters contaminants and chemical tracers in insects: implications for food webs.

Insects are integral to most freshwater and terrestrial food webs, but due to their accumulation of environmental pollutants they are also contaminant vectors that threaten reproduction, development, and survival of consumers. Metamorphosis from larvae to adult can cause large chemical changes in insects, altering contaminant concentrations and fractionation of chemical tracers used to establish contaminant biomagnification in food webs, but no framework exists for predicting and managing these effects. We analyzed data from 39 studies of 68 analytes (stable isotopes and contaminants), and found that metamorphosis effects varied greatly. δ(15)N, widely used to estimate relative trophic position in biomagnification studies, was enriched by ∼ 1‰ during metamorphosis, while δ(13)C used to estimate diet, was similar in larvae and adults. Metals and polycyclic aromatic hydrocarbons (PAHs) were predominantly lost during metamorphosis leading to ∼ 2 to 125-fold higher larval concentrations and higher exposure risks for predators of larvae compared to predators of adults. In contrast, manufactured organic contaminants (such as polychlorinated biphenyls) were retained and concentrated in adults, causing up to ∼ 3-fold higher adult concentrations and higher exposure risks to predators of adult insects. Both food web studies and contaminant management and mitigation strategies need to consider how metamorphosis affects the movement of materials between habitats and ecosystems, with special regard for aquatic-terrestrial linkages.

Unexpected hydrogen isotope variation in oceanic pelagic seabirds.

Hydrogen isotopes have significantly enhanced our understanding of the biogeography of migratory animals. The basis for this methodology lies in predictable, continental patterns of precipitation δD values that are often reflected in an organism's tissues. δD variation is not expected for oceanic pelagic organisms whose dietary hydrogen (water and organic hydrogen in prey) is transferred up the food web from an isotopically homogeneous water source. We report a 142‰ range in the δD values of flight feathers from the Hawaiian petrel (Pterodroma sandwichensis), an oceanic pelagic North Pacific species, and inquire about the source of that variation. We show δD variation between and within four other oceanic pelagic species: Newell's shearwater (Puffinus auricularis newellii), Black-footed albatross (Phoebastria nigripes), Laysan albatross (Phoebastria immutabilis) and Buller's shearwater (Puffinus bulleri). The similarity between muscle δD values of hatch-year Hawaiian petrels and their prey suggests that trophic fractionation does not influence δD values of muscle. We hypothesize that isotopic discrimination is associated with water loss during salt excretion through salt glands. Salt load differs between seabirds that consume isosmotic squid and crustaceans and those that feed on hyposmotic teleost fish. In support of the salt gland hypothesis, we show an inverse relationship between δD and percent teleost fish in diet for three seabird species. Our results demonstrate the utility of δD in the study of oceanic consumers, while also contributing to a better understanding of δD systematics, the basis for one of the most commonly utilized isotope tools in avian ecology.

The early bird gets the shrimp: confronting assumptions of isotopic equilibrium and homogeneity in a wild bird population.

1. Because stable isotope distributions in organic material vary systematically across energy gradients that exist in ecosystems, community and population structures, and in individual physiological systems, isotope values in animal tissues have helped address a broad range of questions in animal ecology. It follows that every tissue sample provides an isotopic profile that can be used to study dietary or movement histories of individual animals. Interpretations of these profiles depend on the assumption that metabolic pools are isotopically well mixed and in equilibrium with dietary resources prior to tissue synthesis, and they extend to the population level by assuming isotope profiles are identically distributed for animals using the same proximal dietary resource. As these assumptions are never fully met, studying structure in the variance of tissue isotope values from wild populations is informative. 2. We studied variation in δ(13) C, δ(15) N, δ(2) H and δ(18) O data for feathers from a population of eared grebes (Podiceps nigricollis) that migrate to Great Salt Lake each fall to moult feathers. During this time, they cannot fly and feed almost exclusively on superabundant brine shrimp (Artemia franciscana). The ecological simplicity of this situation minimized the usual spatial and trophic complexities often present in natural studies of feather isotope values. 3. Ranges and variances of isotope values for the feathers were larger than those from previously published studies that report feather isotopic variance, but they were bimodally distributed in all isotope dimensions. Isotope values for proximal dietary resources and local surface water show that some of the feathers we assumed to have been grown locally must have been grown before birds reached isotopic equilibrium with local diet or immediately prior to arrival at Great Salt Lake. 4. Our study provides novel insights about resource use strategies in eared grebes during migration. More generally, it demonstrates the utility of studying variance structures and questioning assumptions implicit in the interpretation of stable isotope data from wild animals.

Foraging segregation and genetic divergence between geographically proximate colonies of a highly mobile seabird.

Foraging segregation may play an important role in the maintenance of animal diversity, and is a proposed mechanism for promoting genetic divergence within seabird species. However, little information exists regarding its presence among seabird populations. We investigated genetic and foraging divergence between two colonies of endangered Hawaiian petrels (Pterodroma sandwichensis) nesting on the islands of Hawaii and Kauai using the mitochondrial Cytochrome b gene and carbon, nitrogen and hydrogen isotope values (δ(13)C, δ(15)N and δD, respectively) of feathers. Genetic analyses revealed strong differentiation between colonies on Hawaii and Kauai, with Φ(ST) = 0.50 (p < 0.0001). Coalescent-based analyses gave estimates of <1 migration event per 1,000 generations. Hatch-year birds from Kauai had significantly lower δ(13)C and δ(15)N values than those from Hawaii. This is consistent with Kauai birds provisioning chicks with prey derived from near or north of the Hawaiian Islands, and Hawaii birds provisioning young with prey from regions of the equatorial Pacific characterized by elevated δ(15)N values at the food web base. δ(15)N values of Kauai and Hawaii adults differed significantly, indicating additional foraging segregation during molt. Feather δD varied from -69 to 53‰. This variation cannot be related solely to an isotopically homogeneous ocean water source or evaporative water loss. Instead, we propose the involvement of salt gland excretion. Our data demonstrate the presence of foraging segregation between proximately nesting seabird populations, despite high species mobility. This ecological diversity may facilitate population coexistence, and its preservation should be a focus of conservation strategies.

Increased Mercury and Reduced Insect Diversity in Linked Stream-Riparian Food Webs Downstream of a Historical Mercury Mine.

Historical mining left a legacy of abandoned mines and waste rock in remote headwaters of major river systems in the western United States. Understanding the influence of these legacy mines on culturally and ecologically important downstream ecosystems is not always straightforward because of elevated natural levels of mineralization in mining-impacted watersheds. To test the ecological effects of historic mining in the headwaters of the upper Salmon River watershed in Idaho (USA), we measured multiple community and chemical endpoints in downstream linked aquatic-terrestrial food webs. Mining inputs impacted downstream food webs through increased mercury accumulation and decreased insect biodiversity. Total mercury (THg) in seston, aquatic insect larvae, adult aquatic insects, riparian spiders, and fish at sites up to 7.6 km downstream of mining was found at much higher concentrations (1.3-11.3-fold) and was isotopically distinct compared with sites immediately upstream of mining inputs. Methylmercury concentrations in bull trout and riparian spiders were sufficiently high (732-918 and 347-1140 ng MeHg g-1 dry wt, respectively) to affect humans, birds, and piscivorous fish. Furthermore, the alpha-diversity of benthic insects was locally depressed by 12%-20% within 4.3-5.7 km downstream from the mine. However, because total insect biomass was not affected by mine inputs, the mass of mercury in benthic insects at a site (i.e., ng Hg m-2 ) was extremely elevated downstream (10-1778-fold) compared with directly upstream of mining inputs. Downstream adult aquatic insect-mediated fluxes of THg were also high (~16 ng THg m-2 day-1 ). Abandoned mines can have ecologically important effects on downstream communities, including reduced biodiversity and increased mercury flux to higher order consumers, including fish, birds, and humans. Environ Toxicol Chem 2022;41:1696-1710. Published 2022. This article is a U.S. Government work and is in the public domain in the USA.

A migratory divide spanning two continents is associated with genomic and ecological divergence.

Migratory divides are contact zones between breeding populations with divergent migratory strategies during the nonbreeding season. These locations provide an opportunity to evaluate the role of seasonal migration in the maintenance of reproductive isolation, particularly the relationship between population structure and features associated with distinct migratory strategies. We combine light-level geolocators, genomic sequencing, and stable isotopes to investigate the timing of migration and migratory routes of individuals breeding on either side of a migratory divide coinciding with genomic differentiation across a hybrid zone between barn swallow (Hirundo rustica) subspecies in China. Individuals west of the hybrid zone, with H. r. rustica ancestry, had comparatively enriched stable-carbon and hydrogen isotope values and overwintered in eastern Africa, whereas birds east of the hybrid zone, with H. r. gutturalis ancestry, had depleted isotope values and migrated to southern India. The two subspecies took divergent migratory routes around the high-altitude Karakoram Range and arrived on the breeding grounds over 3 weeks apart. These results indicate that assortative mating by timing of arrival and/or selection against hybrids with intermediate migratory traits may maintain reproductive isolation between the subspecies, and that inhospitable geographic features may have contributed to the diversification of Asian avifauna by influencing migratory patterns.

Mercury bioaccumulation and biomagnification in Ozark stream ecosystems.

Crayfish (Orconectes spp.), Asian clam (Corbicula fluminea), northern hog sucker (hog sucker; Hypentelium nigricans), and smallmouth bass (smallmouth; Micropterus dolomieu) from streams in southeastern Missouri (USA) were analyzed for total mercury (HgT) and for stable isotopes of carbon (δ¹³C), nitrogen (δ¹5N), and sulfur (δ³4S) to discern Hg transfer pathways. HgT concentrations were generally lowest in crayfish (0.005-0.112 µg/g dw) and highest in smallmouth (0.093-4.041 µg/g dw), as was δ¹5N. HgT was also lower and δ¹5N was higher in all biota from a stream draining a more heavily populated historical lead-zinc mining area than from similar sites with mostly undeveloped forested watersheds. δ¹³C in biota was lowest at spring-influenced sites, reflecting CO2 inputs and temperature influences, and δ³4S increased from south to north in all taxa. However, HgT was not strongly correlated with either δ¹³C or δ³4S in biota. Trophic position (TP) computed from crayfish δ¹5N was lower in hog suckers (mean=2.8) than in smallmouth (mean=3.2), but not at all sites. HgT, δ¹³C, δ³4S, and TP in hog suckers increased with total length (length) at some sites, indicating site-specific ontogenetic diet shifts. Changes with length were less evident in smallmouth. Length-adjusted HgT site means in both species were strongly correlated with HgT in crayfish (r²=0.97, P<0.01), but not with HgT in Corbicula (r²=0.02, P>0.05). ANCOVA and regression models incorporating only TP and, for hog suckers, length, accurately and precisely predicted HgT concentrations in both fish species from all locations. Although low compared to many areas of the USA, HgT (and therefore methylmercury) concentrations in smallmouth and hog suckers are sufficiently high to represent a threat to human health and wildlife. Our data indicate that in Ozark streams, Hg concentrations in crayfish are at least partly determined by their diet, with concentrations in hog suckers, smallmouth, and possibly other higher-level consumers largely determined by concentrations in crayfish and other primary and secondary consumers, fish growth rates, and TP.

Energetic and health effects of protein overconsumption constrain dietary adaptation in an apex predator.

Studies of predator feeding ecology commonly focus on energy intake. However, captive predators have been documented to selectively feed to optimize macronutrient intake. As many apex predators experience environmental changes that affect prey availability, limitations on selective feeding can affect energetics and health. We estimated the protein:fat ratio of diets consumed by wild polar bears using a novel isotope-based approach, measured protein:fat ratios selected by zoo polar bears offered dietary choice and examined potential energetic and health consequences of overconsuming protein. Dietary protein levels selected by wild and zoo polar bears were low and similar to selection observed in omnivorous brown bears, which reduced energy intake requirements by 70% compared with lean meat diets. Higher-protein diets fed to zoo polar bears during normal care were concurrent with high rates of mortality from kidney disease and liver cancer. Our results suggest that polar bears have low protein requirements and that limitations on selective consumption of marine mammal blubber consequent to climate change could meaningfully increase their energetic costs. Although bear protein requirements appear lower than those of other carnivores, the energetic and health consequences of protein overconsumption identified in this study have the potential to affect a wide range of taxa.