The geographic extent of bird populations affected by renewable-energy development.

Bird populations are declining globally. Wind and solar energy can reduce emissions of fossil fuels that drive anthropogenic climate change, yet renewable-energy production represents a potential threat to bird species. Surveys to assess potential effects at renewable-energy facilities are exclusively local, and the geographic extent encompassed by birds killed at these facilities is largely unknown, which creates challenges for minimizing and mitigating the population-level and cumulative effects of these fatalities. We performed geospatial analyses of stable hydrogen isotope data obtained from feathers of 871 individuals of 24 bird species found dead at solar- and wind-energy facilities in California (USA). Most species had individuals with a mix of origins, ranging from 23% to 98% nonlocal. Mean minimum distances to areas of likely origin for nonlocal individuals were as close as 97 to >1250 km, and these minimum distances were larger for species found at solar-energy facilities in deserts than at wind-energy facilities in grasslands (Cohen's d = 6.5). Fatalities were drawn from an estimated 30-100% of species' desingated ranges, and this percentage was significantly smaller for species with large ranges found at wind facilities (Pearson's r = -0.67). Temporal patterns in the geographic origin of fatalities suggested that migratory movements and nonmigratory movements, such as dispersal and nomadism, influence exposure to fatality risk for these birds. Our results illustrate the power of using stable isotope data to assess the geographic extent of renewable-energy fatalities on birds. As the buildout of renewable-energy facilities continues, accurate assessment of the geographic footprint of wildlife fatalities can be used to inform compensatory mitigation for their population-level and cumulative effects.

Extensión geográfica de las poblaciones de aves afectadas por desarrollos de energía renovable Resumen Las poblaciones mundiales de aves están en declive. Las energías solar y eólica pueden reducir las emisiones de combustibles fósiles que causan el cambio climático, aunque la producción de energías renovables representa una amenaza potencial para las aves. Los censos para evaluar los efectos potenciales en los centros de energía renovable son exclusivamente locales y se sabe poco sobre la extensión geográfica representada por las aves que mueren en estas instalaciones, lo que plantea obstáculos para mitigar los efectos acumulativos y de nivel poblacional de estas muertes. Realizamos análisis geoespaciales con datos del isótopo de hidrógeno estable obtenido de las plumas de 871 ejemplares de 24 especies de aves que fueron hallados muertos en los centros de energía solar y eólica en California, EE.UU. La mayoría de las especies contó con ejemplares de orígenes mixtos, con un rango del 23% al 98% no local. La media de la distancia mínima a las áreas de probable origen de los ejemplares no locales varía entre los 97 hasta > 1,250 km. Estas distancias mínimas fueron mayores para las especies encontradas en los centros de energía solar situadas en desiertos que para las especies encontradas en los centros de energía eólica localizadas en pastizales (d de Cohen = 6.5). Las muertes representan un 30­100% de la extensión de las especies. Este porcentaje fue significativamente menor para las especies con extensiones amplias encontradas en instalaciones eólicas (r de Pearson = ­0.67). Los patrones temporales en el origen geográfico de las muertes sugieren que los movimientos migratorios y no migratorios, como la dispersión y el nomadismo, influyen en la exposición de estas aves al riesgo de muerte. Nuestros resultados demuestran la utilidad de los isótopos estables para evaluar el alcance geográfico de las muertes de aves asociadas a energías renovables. Con el progresivo aumento de instalaciones de energía renovable, una evaluación precisa de la huella geográfica de la mortandad de fauna salvaje podrá guiar la mitigación compensatoria de sus efectos acumulativos y de nivel poblacional.

Black Bear Behavior and Movements Are Not Definitive Measures of Anthropogenic Food Use.

Increasing human-bear conflicts are a growing concern, and managers often assume bears in developed areas are food-conditioned. We examined the relationship between human-bear conflicts and food conditioning by analyzing isotopic values of hair from black bears (Ursus americanus floridanus) involved in research (n = 34) and conflicts (n = 45). We separated research bears into wild and developed subgroups based on the impervious surface within their home ranges and separated conflict bears based on observations of human food consumption (anthropogenic = observations; management = no observations). We initially assumed wild bears were not food conditioned and anthropogenic bears were. However, using isotopic values, we classified 79% of anthropogenic bears and 8% of wild bears as food conditioned. Next, we assigned these bears to the appropriate food conditioned category and used the categorizations as a training set to classify developed and management bears. We estimated that 53% of management bears and 20% of developed bears were food conditioned. Only 60% of bears captured within or using developed areas showed evidence of food conditioning. We also found that δ13C values were a better predictor of anthropogenic foods in a bear's diet than δ15N values. Our results indicate that bears in developed areas are not necessarily food conditioned and caution against management actions based on limited observations of bear behavior.

Long-term stability in the circumpolar foraging range of a Southern Ocean predator between the eras of whaling and rapid climate change.

Assessing environmental changes in Southern Ocean ecosystems is difficult due to its remoteness and data sparsity. Monitoring marine predators that respond rapidly to environmental variation may enable us to track anthropogenic effects on ecosystems. Yet, many long-term datasets of marine predators are incomplete because they are spatially constrained and/or track ecosystems already modified by industrial fishing and whaling in the latter half of the 20th century. Here, we assess the contemporary offshore distribution of a wide-ranging marine predator, the southern right whale (SRW, Eubalaena australis), that forages on copepods and krill from ~30°S to the Antarctic ice edge (>60°S). We analyzed carbon and nitrogen isotope values of 1,002 skin samples from six genetically distinct SRW populations using a customized assignment approach that accounts for temporal and spatial variation in the Southern Ocean phytoplankton isoscape. Over the past three decades, SRWs increased their use of mid-latitude foraging grounds in the south Atlantic and southwest (SW) Indian oceans in the late austral summer and autumn and slightly increased their use of high-latitude (>60°S) foraging grounds in the SW Pacific, coincident with observed changes in prey distribution and abundance on a circumpolar scale. Comparing foraging assignments with whaling records since the 18th century showed remarkable stability in use of mid-latitude foraging areas. We attribute this consistency across four centuries to the physical stability of ocean fronts and resulting productivity in mid-latitude ecosystems of the Southern Ocean compared with polar regions that may be more influenced by recent climate change.

White-Nose Syndrome Pathogen Pseudogymnoascus destructans Detected in Migratory Tree-Roosting Bats.

White-nose syndrome (WNS) is an emerging fungal epizootic disease that has caused large-scale mortality in several species of North American bats. The fungus that causes WNS, Pseudogymnoascus destructans (Pd), has also been detected in bat species without diagnostic signs of WNS. Although these species could play a role in WNS spread, understanding of the spatial and temporal extents of Pd occurrence on WNS-resistant species is limited. This study evaluated the presence of Pd on 272 individuals of three species of migratory tree-roosting bats: hoary (Lasiurus cinereus), eastern red (Lasiurus borealis), and silver-haired (Lasionycteris noctivagans) bats, obtained opportunistically during summer and autumn from throughout much of their ranges in North America. We also compared tissue sampling protocols (i.e., tissue swabbing, fur swabbing, and DNA extraction of excised wing tissue). We detected Pd on three eastern red bats from Illinois and Ohio, US, one silver-haired bat from West Virginia, US, and one hoary bat from New York, US, all via DNA extracted from wing tissue of carcasses. These results document the first publicly reported detections of Pd on a hoary bat and on migratory bats during the autumn migratory period, and demonstrate the potential for using carcasses salvaged at wind-energy facilities to monitor for Pd.

Vulnerability of avian populations to renewable energy production.

Renewable energy production can kill individual birds, but little is known about how it affects avian populations. We assessed the vulnerability of populations for 23 priority bird species killed at wind and solar facilities in California, USA. Bayesian hierarchical models suggested that 48% of these species were vulnerable to population-level effects from added fatalities caused by renewables and other sources. Effects of renewables extended far beyond the location of energy production to impact bird populations in distant regions across continental migration networks. Populations of species associated with grasslands where turbines were located were most vulnerable to wind. Populations of nocturnal migrant species were most vulnerable to solar, despite not typically being associated with deserts where the solar facilities we evaluated were located. Our findings indicate that addressing declines of North American bird populations requires consideration of the effects of renewables and other anthropogenic threats on both nearby and distant populations of vulnerable species.

Hydrogen isotope assimilation and discrimination in green turtles.

Although hydrogen isotopes (δ2H) are commonly used as tracers of animal movement, minimal research has investigated the use of δ2H as a proxy to quantify resource and habitat use. While carbon and nitrogen are ultimately derived from a single source (food), the proportion of hydrogen in consumer tissues originates from two distinct sources: body water and food. Before hydrogen isotopes can be effectively used as a resource and habitat tracer, we need estimates of (net) discrimination factors (Δ2HNet) that account for the physiologically mediated differences in the δ2H values of animal tissues relative to that of the food and water sources they use to synthesize tissues. Here, we estimated Δ2HNet in captive green turtles (Chelonia mydas) by measuring the δ2H values of tissues (epidermis and blood components) and dietary macromolecules collected in two controlled feeding experiments. Tissue δ2H and Δ2HNet values varied systematically among tissues, with epidermis having higher δ2H and Δ2HNet values than blood components, which mirrors patterns between keratinaceous tissues (feathers, hair) and blood in birds and mammals. Serum/plasma of adult female green turtles had significantly lower δ2H values compared with juveniles, likely due to increased lipid mobilization associated with reproduction. This is the first study to quantify Δ2HNet values in a marine ectotherm, and we anticipate that our results will further refine the use of δ2H analysis to better understand animal resource and habitat use in marine ecosystems, especially coastal areas fueled by a combination of marine (e.g. micro/macroalgae and seagrass) and terrestrial (e.g. mangroves) primary production.

Identifying patterns in foraging-area origins in breeding aggregations of migratory species: Loggerhead turtles in the Northwest Atlantic.

Population assessments conducted at reproductive sites of migratory species necessitate understanding the foraging-area origins of breeding individuals. Without this information, efforts to contextualize changes in breeding populations and develop effective management strategies are compromised. We used stable isotope analysis of tissue samples collected from loggerhead sea turtles (Caretta caretta) nesting at seven sites in the Northern Recovery Unit (NRU) of the eastern United States (North Carolina, South Carolina and Georgia) to assign females to three separate foraging areas in the Northwest Atlantic Ocean (NWA). We found that the majority of the females at NRU nesting sites (84.4%) use more northern foraging areas in the Mid-Atlantic Bight, while fewer females use more proximate foraging areas in the South Atlantic Bight (13.4%) and more southerly foraging areas in the Subtropical Northwest Atlantic (2.2%). We did not find significant latitudinal or temporal trends in the proportions of NRU females originating from different foraging areas. Combining these findings with previous data from stable isotope and satellite tracking studies across NWA nesting sites showed that variation in the proportion of adult loggerheads originating from different foraging areas is primarily related differences between recovery units: individuals in the NRU primarily use the Mid-Atlantic Bight foraging area, while individuals from the three Florida recovery units primarily use the Subtropical Northwest Atlantic and Eastern Gulf of Mexico foraging areas. Because each foraging area is associated with its own distinct ecological characteristics, environmental fluctuations and anthropogenic threats that affect the abundance and productivity of individuals at nesting sites, this information is critical for accurately evaluating population trends and developing effective region-specific management strategies.

Mechanistic model predicts tissue-environment relationships and trophic shifts in animal hydrogen and oxygen isotope ratios.

Statistical regression relationships between the hydrogen (H) and oxygen (O) isotope ratios (δ2H and δ18O, respectively) of animal organic tissues and those of environmental water have been widely used to reconstruct animal movements, paleoenvironments, and diet and trophic relationships. In natural populations, however, tissue-environment isotopic relationships are highly variable among animal types and geographic regions. No systematic understanding of the origin(s) of this variability currently exists, clouding the interpretation of isotope data. Here, we present and apply a model, based on fundamental metabolic relationships, to test the sensitivity of consumer tissue H and O isotope ratios, and thus tissue-environment relationships, to basic physiological, behavioral, and environmental parameters. We then simulate patterns in consumer tissue isotopic compositions under several 'real-world' scenarios, demonstrating that the new model can reproduce-and potentially explain-previously observed patterns in consumer tissue H isotope ratios, including between-continent differences in feather-precipitation relationships and 2H-enrichment with trophic level across species. The model makes several fundamental predictions about the organic O isotope system, which constitute hypotheses for future testing as new data are obtained. By highlighting potential sources of variability and bias in tissue-environment relationships and establishing a framework within which such effects can be predicted, these results should advance the application of H and O isotopes in ecological, paleoecological, and forensic research.

Effect of heat and singeing on stable hydrogen isotope ratios of bird feathers and implications for their use in determining geographic origin.

RATIONALE: Stable hydrogen isotope (δ2 H) ratios of animal tissues are useful for assessing movement and geographic origin of mobile organisms. However, it is uncertain whether heat and singeing affects feather δ2 H values and thus subsequent geographic assignments. This is relevant for birds of conservation interest that are burned and killed at concentrating solar-energy facilities that reflect sunlight to a receiving tower and generate a solar flux field. METHODS: We used a controlled experiment to test the effect of known heat loads (exposure to 200, 250 or 300°C for 1 min) on the morphology and δ2 H values of feathers from two songbird species. Subsequently, we examined the effects of singeing on δ2 H values of feathers from three other songbird species that were found dead in the field at a concentrating solar-energy facility. RESULTS: Relative to control samples, heating caused visual morphological changes to feathers, including shriveling at 250°C and charring at 300°C. The δ2 H values significantly declined by a mean of 27.8‰ in experimental samples exposed to 300°C. There was no statistically detectable difference between δ2 H values of the singed and unsinged portions of field-collected feathers from the same bird. CONCLUSIONS: Limited singeing that did not dramatically alter the feather morphology did not substantially affect δ2 H values of feathers from these songbirds. However, higher temperatures induced charring and reduced δ2 H values. Therefore, severely charred feathers should be avoided when selecting feathers for δ2 H-based assessment of geographic origin.

Biomarkers reveal sea turtles remained in oiled areas following the Deepwater Horizon oil spill.

Assessments of large-scale disasters, such as the Deepwater Horizon oil spill, are problematic because while measurements of post-disturbance conditions are common, measurements of pre-disturbance baselines are only rarely available. Without adequate observations of pre-disaster organismal and environmental conditions, it is impossible to assess the impact of such catastrophes on animal populations and ecological communities. Here, we use long-term biological tissue records to provide pre-disaster data for a vulnerable marine organism. Keratin samples from the carapace of loggerhead sea turtles record the foraging history for up to 18 years, allowing us to evaluate the effect of the oil spill on sea turtle foraging patterns. Samples were collected from 76 satellite-tracked adult loggerheads in 2011 and 2012, approximately one to two years after the spill. Of the 10 individuals that foraged in areas exposed to surface oil, none demonstrated significant changes in foraging patterns post spill. The observed long-term fidelity to foraging sites indicates that loggerheads in the northern Gulf of Mexico likely remained in established foraging sites, regardless of the introduction of oil and chemical dispersants. More research is needed to address potential long-term health consequences to turtles in this region. Mobile marine organisms present challenges for researchers to monitor effects of environmental disasters, both spatially and temporally. We demonstrate that biological tissues can reveal long-term histories of animal behavior and provide critical pre-disaster baselines following an anthropogenic disturbance or natural disaster.

Stable isotopic comparison between loggerhead sea turtle tissues.

RATIONALE: Stable isotope analysis has been used extensively to provide ecological information about diet and foraging location of many species. The difference in isotopic composition between animal tissue and its diet, or the diet-tissue discrimination factor, varies with tissue type. Therefore, direct comparisons between isotopic values of tissues are inaccurate without an appropriate conversion factor. We focus on the loggerhead sea turtle (Caretta caretta), for which a variety of tissues have been used to examine diet, habitat use, and migratory origin through stable isotope analysis. We calculated tissue-to-tissue conversions between two commonly sampled tissues. METHODS: Epidermis and scute (the keratin covering on the carapace) were sampled from 33 adult loggerheads nesting at two beaches in Florida (Casey Key and Canaveral National Seashore). Carbon and nitrogen stable isotope ratios were measured in the epidermis and the youngest portion of the scute tissue, which reflect the isotopic composition of the diet and habitat over similar time periods of the order of several months. RESULTS: Significant linear relationships were observed between the δ(13)C and δ(15)N values of these two tissues, indicating they can be converted reliably. CONCLUSIONS: Whereas both epidermis and scute samples are commonly sampled from nesting sea turtles to study trophic ecology and habitat use, the data from these studies have not been comparable without reliable tissue-to-tissue conversions. The equations provided here allow isotopic datasets using the two tissues to be combined in previously published and subsequent studies of sea turtle foraging ecology and migratory movement. In addition, we recommend that future isotopic comparisons between tissues of any organism utilize linear regressions to calculate tissue-to-tissue conversions.

Mother-egg stable isotope conversions and effects of lipid extraction and ethanol preservation on loggerhead eggs.

Carbon and nitrogen stable isotope (δ(13)C and δ(15)N) analysis has been used to elucidate foraging and migration behaviours of endangered sea turtle populations. Isotopic analysis of tissue samples from nesting females can provide information about their foraging locations before reproduction. To determine whether loggerhead (Caretta caretta) eggs provide a good proxy for maternal isotope values, we addressed the following three objectives: (i) we evaluated isotopic effects of ethanol preservation and lipid extraction on yolk; (ii) we examined the isotopic offset between maternal epidermis and corresponding egg yolk and albumen tissue δ(13)C and δ(15)N values; and (iii) we assessed the accuracy of foraging ground assignment using egg yolk and albumen stable isotope values as a proxy for maternal epidermis. Epidermis (n = 61), albumen (n = 61) and yolk samples (n = 24) were collected in 2011 from nesting females at Wassaw Island, GA, USA. Subsamples from frozen and ethanol-preserved yolk samples were lipid extracted. Both lipid extraction and ethanol preservation significantly affected yolk δ(13)C, while δ(15)N values were not altered at a biologically relevant level. The mathematical corrections provided here allow for normalization of yolk δ(13)C values with these treatments. Significant tissue conversion equations were found between δ(13)C and δ(15)N values of maternal epidermis and corresponding yolk and albumen. Finally, the consistency in assignment to a foraging area was high (up to 84%), indicating that these conversion equations can be used in future studies where stable isotopes are measured to determine female foraging behaviour and trophic relationships by assessing egg components. Loggerhead eggs can thus provide reliable isotopic information when samples from nesting females cannot be obtained.

Temporal consistency and individual specialization in resource use by green turtles in successive life stages.

Not all individuals in a population use the same subset of dietary and habitat resources. Patterns of individual specialization have been documented in an increasing number of organisms, but often without an associated time scale over which niche specialization was observed. We examined the patterns in individual resource use through time and in relation to the population with metrics of temporal consistency and degree of individual specialization. We used stable isotope analysis of carbon and nitrogen in successive subsections of scute tissue from the carapace to compare foraging patterns in three successive life stages of the green turtle (Chelonia mydas). Temporal consistency was measured as the mean within-individual variation in stable isotope values through time, whereas the degree of individual specialization was a ratio of the individual variation to that of the population. The distinction between these two parameters is important, as the metric of temporal consistency quantifies the regularity of individual resource use, and the degree of individual specialization indicates what proportion of the population niche an average individual uses. The scute record retains a chronological history of resource use and was estimated to represent a minimum 0.8 years in juveniles to a maximum of 6.5 years in adults. Both temporal consistency and individual specialization varied significantly among life stages. Adults were highly consistent in resource use through time and formed a generalist population with individual specialists maintaining long-term patterns in resource use. Oceanic and neritic juvenile life stages exhibited less temporal consistency in resource use with less individual specialization than adults. These observations are important when considering the ecological roles filled by green turtles in each life stage; also, individual differences in resource use may result in differential fitness consequences.

Inherent variation in stable isotope values and discrimination factors in two life stages of green turtles.

We examine inherent variation in carbon and nitrogen stable isotope values of multiple soft tissues from a population of captive green turtles Chelonia mydas to determine the extent of isotopic variation due to individual differences in physiology. We compare the measured inherent variation in the captive population with the isotopic variation observed in a wild population of juvenile green turtles. Additionally, we measure diet-tissue discrimination factors to determine the offset that occurs between isotope values of the food source and four green turtle tissues. Tissue samples (epidermis, dermis, serum, and red blood cells) were collected from captive green turtles in two life stages (40 large juveniles and 30 adults) at the Cayman Turtle Farm, Grand Cayman, and analyzed for carbon and nitrogen stable isotopes. Multivariate normal models were fit to the isotope data, and the Bayesian Information Criterion was used for model selection. Inherent variation and discrimination factors differed among tissues and life stages. Inherent variation was found to make up a small portion of the isotopic variation measured in a wild population. Discrimination factors not only are tissue and life stage dependent but also appear to vary with diet and sea turtle species, thus highlighting the need for appropriate discrimination factors in dietary reconstructions and trophic-level estimations. Our measures of inherent variation will also be informative in field studies employing stable isotope analysis so that differences in diet or habitat are more accurately identified.

Individual specialists in a generalist population: results from a long-term stable isotope series.

Individual variation in resource use has often been ignored in ecological studies, but closer examination of individual patterns through time may reveal significant intrapopulation differences. Adult loggerhead sea turtles (Caretta caretta) are generalist carnivores with a wide geographical range, resulting in a broad isotopic niche. We microsampled scute, a persistent and continuously growing tissue, to examine long-term variation in resource use (up to 12 years) in 15 nesting loggerhead turtles. Using stable isotopes of nitrogen and carbon, we examined the resource use patterns (integration of diet, habitat and geographical location) and demonstrate that individual loggerheads are long-term specialists within a generalist population. We present our results in the context of a conceptual model comparing isotopic niches in specialist and generalist populations. Individual consistency may have important ecological, evolutionary and conservation consequences, such as the reduction of intraspecific competition.

Energetic basis of colonial living in social insects.

Understanding the ecology and evolution of insect societies requires greater knowledge of how sociality affects the performance of whole colonies. Metabolic scaling theory, based largely on the body mass scaling of metabolic rate, has successfully predicted many aspects of the physiology and life history of individual (or unitary) organisms. Here we show, using a diverse set of social insect species, that this same theory predicts the size dependence of basic features of the physiology (i.e., metabolic rate, reproductive allocation) and life history (i.e., survival, growth, and reproduction) of whole colonies. The similarity in the size dependence of these features in unitary organisms and whole colonies points to commonalities in functional organization. Thus, it raises an important question of how such evolutionary convergence could arise through the process of natural selection.