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Analysis of annual dissolved-solids loading from selected natural and irrigated catchments in the Upper Colorado River Basin, 1974-2003

Dissolved-solids loading from 17 natural catchments and 14 irrigated catchments in the Upper Colorado River Basin was examined for the period from 1974 through 2003. In general, dissolved-solids loading increased and decreased concurrently in natural and irrigated catchments but at different magnitudes. Annually, the magnitude of loading in natural catchments changed about 10 percent more, on average, than in irrigated catchments. Measures of variability, or spread, indicate that natural catchments had 35 percent greater annual variability in loading than irrigated catchments. Precipitation and dissolved-solids loads were positively correlated in natural catchments, and a weak positive correlation was determined for irrigated catchments. A weak negative correlation between temperature and dissolved-solids load was determined for both natural and irrigated catchments. In irrigated catchments, the dissolved-solids load response to an above-average precipitation period from 1982 through 1987 generally lagged behind that in the natural catchments. On average, irrigated catchments with reservoir storage had the largest normalized maximum annual loads during the wet period.

Arizona, Colorado, New Mexico, Utah, Wyoming

Genomics of Arctic cod

The Arctic cod (Boreogadus saida) is an abundant marine fish that plays a vital role in the marine food web. To better understand the population genetic structure and the role of natural selection acting on the maternally-inherited mitochondrial genome (mitogenome), a molecule often associated with adaptations to temperature, we analyzed genetic data collected from 11 biparentally-inherited nuclear microsatellite DNA loci and nucleotide sequence data from from the mitochondrial DNA (mtDNA) cytochrome b (cytb) gene and, for a subset of individuals, the entire mitogenome. In addition, due to potential of species misidentification with morphologically similar Polar cod (Arctogadus glacialis), we used ddRAD-Seq data to determine the level of divergence between species and identify species-specific markers. Based on the findings presented here, Arctic cod across the Pacific Arctic (Bering, Chukchi, and Beaufort Seas) comprise a single panmictic population with high genetic diversity compared to other gadids. High genetic diversity was indicated across all 13 protein-coding genes in the mitogenome. In addition, we found moderate levels of genetic diversity in the nuclear microsatellite loci, with highest diversity found in the Chukchi Sea. Our analyses of markers from both marker classes (nuclear microsatellite fragment data and mtDNA cytb sequence data) failed to uncover a signal of microgeographic genetic structure within Arctic cod across the three regions, within the Alaskan Beaufort Sea, or between near-shore or offshore habitats. Further, data from a subset of mitogenomes revealed no genetic differentiation between Bering, Chukchi, and Beaufort seas populations for Arctic cod, Saffron cod (Eleginus gracilis), or Walleye pollock (Gadus chalcogrammus). However, we uncovered significant differences in the distribution of microsatellite alleles between the southern Chukchi and central and eastern Beaufort Sea samples of Arctic cod. Finally, using ddRAD-Seq data, we identified species-specific markers and in conjunction with mitogenome data, identified an Arctic cod x Polar cod hybrid in western Canadian Beaufort Sea. Overall, the lack of genetic structure among Arctic cod within the Bering, Chukchi and Beaufort seas of Alaska is concordant with the absence of geographic barriers to dispersal and typical among marine fishes. Arctic cod may exhibit a genetic pattern of isolation-by-distance, whereby populations in closer geographic proximity are more genetically similar than more distant populations. As this signal is only found between our two fartherest localities, data from populations elsewhere in the species’ global range are needed to determine if this is a general characteristic. Further, tests for selection suggested a limited role for natural selection acting on the mitochondrial genome of Arctic cod, but do not exclude the possibility of selection on genes involved in nuclear-mitogenome interactions. Unlike previous genetic assessment of Arctic cod sampled from the Chukchi Sea, the high levels of genetic diversity found in Arctic cod assayed in this study, across regions, suggests that the species in the Beaufort and Chukchi seas does not suffer from low levels of genetic variation, at least at neutral genetic markers. The large census size of Arctic cod may allow this species to retain high levels of genetic diversity. In addition, we discovered the presence of hybridization between Arctic and Polar cod (although low in frequency). Hybridization is expected to occur when environmental changes modify species distributions that result in contact between species that were previously separated. In such cases, hybridization may be an evolutionary mechanism that promotes an increase in genetic diversity that may provide species occupying changing environments with locally-adapted genotypes and, therefore, phenotypes. Natural selection can only act on the standing genetic variation present within a population. Therefore, given its higher levels of genetic diversity in combination with a large population size, Arctic cod may be resilient to current and future environmental change, as high genetic diversity is expected to increase opportunities for positive selection to act on genetic variants beneficial in different environments, regardless of the source of that genetic variation.

OCS Study

Consequences of least tern ( Sternula antillarum ) microhabitat nest-site selection on natural and mechanically constructed sandbars in the Missouri River

Nest-habitat selection in colonial species has rarely been assessed at multiple spatial scales to evaluate its fitness consequences. Management for the federally endangered U.S. Interior population of Least Terns (Sternula antillarum) has focused on maintenance of breeding habitats, including mechanical construction of sandbars from dredged material. Least Terns are attracted to large areas of unvegetated substrate, yet small-scale habitat features are thought to trigger selection for nesting. We evaluated nest-scale habitat selection to determine (1) whether selection differs between constructed and natural sandbars and (2) the subsequent consequences of habitat selection on nest success. During 2006–2008, we examined 869 Least Tern nest sites on constructed and natural sandbars in the Missouri River for evidence of microhabitat selection at the nest in relation to habitat within the surrounding 3-m area. Least Tern nest sites had coarser and larger substrate materials at the nest, more debris, and less vegetation than the surrounding area. Nests in constructed habitats had a greater percentage of coarse substrates and less vegetation or debris than nests in naturally created habitats. Apparent nest success was 1.8× greater on constructed than on natural sandbars. Nest success was best predicted by models with two spatial scales of predictors, including substrates (nest) and vegetation and debris (nest or surrounding area). Our results indicate that Least Terns select nest microhabitat characteristics that are associated with wind- and water-scoured habitats, and that nest success increases when these habitats are selected.

Nebraska;South Dakota

Occam's shadow: levels of analysis in evolutionary ecology - where to next?

Evolutionary ecology is the study of evolutionary processes, and the ecological conditions that influence them. A fundamental paradigm underlying the study of evolution is natural selection. Although there are a variety of operational definitions for natural selection in the literature, perhaps the most general one is that which characterizes selection as the process whereby heritable variation in fitness associated with variation in one or more phenotypic traits leads to intergenerational change in the frequency distribution of those traits. The past 20 years have witnessed a marked increase in the precision and reliability of our ability to estimate one or more components of fitness and characterize natural selection in wild populations, owing particularly to significant advances in methods for analysis of data from marked individuals. In this paper, we focus on several issues that we believe are important considerations for the application and development of these methods in the context of addressing questions in evolutionary ecology. First, our traditional approach to estimation often rests upon analysis of aggregates of individuals, which in the wild may reflect increasingly non-random (selected) samples with respect to the trait(s) of interest. In some cases, analysis at the aggregate level, rather than the individual level, may obscure important patterns. While there are a growing number of analytical tools available to estimate parameters at the individual level, and which can cope (to varying degrees) with progressive selection of the sample, the advent of new methods does not reduce the need to consider carefully the appropriate level of analysis in the first place. Estimation should be motivated a priori by strong theoretical analysis. Doing so provides clear guidance, in terms of both (i) assisting in the identification of realistic and meaningful models to include in the candidate model set, and (ii) providing the appropriate context under which the results are interpreted. Second, while it is true that selection (as defined) operates at the level of the individual, the selection gradient is often (if not generally) conditional on the abundance of the population. As such, it may be important to consider estimating transition rates conditional on both the parameter values of the other individuals in the population (or at least their distribution), and population abundance. This will undoubtedly pose a considerable challenge, for both single- and multi-strata applications. It will also require renewed consideration of the estimation of abundance, especially for open populations. Thirdly, selection typically operates on dynamic, individually varying traits. Such estimation may require characterizing fitness in terms of individual plasticity in one or more state variables, constituting analysis of the norms of reaction of individuals to variable environments. This can be quite complex, especially for traits that are under facultative control. Recent work has indicated that the pattern of selection on such traits is conditional on the relative rates of movement among and frequency of spatially heterogeneous habitats, suggesting analyses of evolution of life histories in open populations can be misleading in some cases.

Journal of Applied Statistics

How organisms do the right thing: The attractor hypothesis

Neo-Darwinian theory is highly successful at explaining the emergence of adaptive traits over successive generations. However, there are reasons to doubt its efficacy in explaining the observed, impressively detailed adaptive responses of organisms to day-to-day changes in their surroundings. Also, the theory lacks a clear mechanism to account for both plasticity and canalization. In effect, there is a growing sentiment that the neo-Darwinian paradigm is incomplete, that something more than genetic structure, mutation, genetic drift, and the action of natural selection is required to explain organismal behavior. In this paper we extend the view of organisms as complex self-organizing entities by arguing that basic physical laws, coupled with the acquisitive nature of organisms, makes adaptation all but tautological. That is, much adaptation is an unavoidable emergent property of organisms' complexity and, to some a significant degree, occurs quite independently of genomic changes wrought by natural selection. For reasons that will become obvious, we refer to this assertion as the attractor hypothesis. The arguments also clarify the concept of "adaptation." Adaptation across generations, by natural selection, equates to the (game theoretic) maximization of fitness (the success with which one individual produces more individuals), while self-organizing based adaptation, within generations, equates to energetic efficiency and the matching of intake and biosynthesis to need. Finally, we discuss implications of the attractor hypothesis for a wide variety of genetical and physiological phenomena, including genetic architecture, directed mutation, genetic imprinting, paramutation, hormesis, plasticity, optimality theory, genotype-phenotype linkage and puncuated equilibrium, and present suggestions for tests of the hypothesis. ?? 1998 American Institute of Physics.

Chaos

Adaptive population divergence and directional gene flow across steep elevational gradients in a climate‐sensitive mammal

The American pika is a thermally sensitive, alpine lagomorph species. Recent climate-associated population extirpations and genetic signatures of reduced population sizes range-wide indicate the viability of this species is sensitive to climate change. To test for potential adaptive responses to climate stress, we sampled pikas along two elevational gradients (each ~470 to 1640 m) and employed three outlier detection methods, BAYESCAN, LFMM, and BAYPASS, to scan for genotype-environment associations in samples genotyped at 30,763 SNP loci. We resolved 173 loci with robust evidence of natural selection detected by either two independent analyses or replicated in both transects. A BLASTN search of these outlier loci revealed several genes associated with metabolic function and oxygen transport, indicating natural selection from thermal stress and hypoxia. We also found evidence of directional gene flow primarily downslope from large high-elevation populations and reduced gene flow at outlier loci, a pattern suggesting potential impediments to the upward elevational movement of adaptive alleles in response to contemporary climate change. Finally, we documented evidence of reduced genetic diversity associated the south-facing transect and an increase in corticosterone stress levels associated with inbreeding. This study suggests the American pika is already undergoing climate-associated natural selection at multiple genomic regions. Further analysis is needed to determine if the rate of climate adaptation in the American pika and other thermally sensitive species will be able to keep pace with rapidly changing climate conditions.

Washington

Quantity, structure, and habitat selection of natural spawning reefs by walleyes in a north temperate lake: A multiscale analysis

Spawning habitat, the cornerstone of self-sustaining, naturally reproducing walleye Sander vitreus populations, has received limited quantitative research. Our goal was to quantitatively describe the structure and quantity of natural walleye spawning habitat and evaluate potential selection of habitat in Big Crooked Lake, Wisconsin. In 2004 and 2005, we located and delineated walleye egg deposition polygons through visual snorkel and scuba surveys. We also delineated recently deposited, adhesive egg patches daily along one spawning reef in 2005. To determine habitat selection, we quantified and compared spawning and lakewide available habitat at different scales. In both years, walleyes used similar spawning habitat, including three geomorphic types: linear shorelines, a point bar, and an island. Walleyes used only 14% of the entire lake shoreline and 39% of the shoreline comprised of gravel (6.4–76.0 mm), cobble (76.1–149.9 mm), or coarser substrates for spawning in 2005, indicating selection of specific spawning habitat. Lakewide, walleyes spawned close to shore (outer egg deposition polygon boundary mean distance = 2.7 m), in shallow water (outer egg deposition polygon boundary mean depth = 0.3 m), and over gravel substrate (percent coverage mean = 64.3) having low embeddedness (mean = 1.30). Our best nearshore (0–13-m) resource selection function predicted an increase in the relative probability of egg deposition with the increasing abundance of gravel, cobble, and rubble (150.0–303.9-mm) substrates and a decrease with increasing distance from shore and water depth (89.9% overall correct classification). Adhesive egg patches confirmed that walleyes actively chose nearshore, shallow-water, and coarse-substrate spawning habitat. The quantitative habitat information and predictive models will assist biologists in developing walleye spawning reef protection strategies and potentially aid in designing and evaluating artificial spawning reefs.

Wisconsin

Survival of female Lesser Scaup: Effects of body size, age, and reproductive effort

In birds, larger females generally have greater breeding propensity, reproductive investment, and success than do smaller females. However, optimal female body size also depends on how natural selection acts during other parts of the life cycle. Larger female Lesser Scaup (Aythya affinis) produce larger eggs than do smaller females, and ducklings from larger eggs survive better than those hatching from smaller eggs. Accordingly, we examined patterns of apparent annual survival for female scaup and tested whether natural selection on female body size primarily was stabilizing, a frequent assumption in studies of sexually dimorphic species in which males are the larger sex, or was directional, counter-acting reproductive advantages of large size . We estimated survival using mark-recapture methods for individually marked females from two study sites in Canada (Erickson, Manitoba; St. Denis, Saskatchewan). Structurally larger (adults) and heavier (ducklings) females had lower survival than did smaller individuals in Manitoba; no relationship was detected in adults from Saskatchewan. Survival of adult females declined with indices of increasing reproductive effort at both sites; consequently, the cost of reproduction could explain age -related patterns of breeding propensity in scaup . Furthermore, if larger females are more likely to breed than are smaller females, then cost of reproduction also may help explain why survival was lower for larger females. Overall, we found that advantages of large body size of female scaup during breeding or as young ducklings apparently were counteracted by natural selection favoring lightweight juveniles and structurally smaller adult females through higher annual survival .

Condor

Groundwater quality of the Gulf Coast aquifer system, Houston, Texas, 2010

During March–December 2010, the U.S. Geological Survey, in cooperation with the city of Houston, collected source-water samples from 60 municipal supply wells in the Houston area. These data were collected as part of an ongoing study to determine concentrations, spatial extent, and associated geochemical conditions that might be conducive for mobility and transport of selected naturally occurring contaminants (selected trace elements and radionuclides) in the Gulf Coast aquifer system in the Houston area. In the summers of 2007 and 2008, a reconnaissance-level survey of these constituents in untreated water from 28 municipal supply wells was completed in the Houston area. Included in this report are the complete analytical results for 47 of the 60 samples collected in 2010—those results which were received from the laboratories and reviewed by the authors as of December 31, 2010. All of the wells sampled were screened in the Gulf Coast aquifer system; 22 were screened entirely in the Evangeline aquifer, and the remaining 25 wells contained screened intervals that intersected both Evangeline and Chicot aquifers. The data documented in this report were collected as part of an ongoing study to characterize source-water-quality conditions in untreated groundwater prior to drinking-water treatment. An evaluation of contaminant occurrence in source water provides background information regarding the presence of a contaminant in the environment. Because source-water samples were collected prior to any treatment or blending that potentially could alter contaminant concentrations, the water-quality results documented by this report represent the quality of the source water, not the quality of finished drinking water provided to the public. Samples were analyzed for major ions (calcium, magnesium, potassium, sodium, bromide, chloride, fluoride, silica, and sulfate), residue on evaporation (dissolved solids), trace elements (arsenic, barium, boron, chromium, iron, lithium, manganese, molybdenum, selenium, strontium, and vanadium), and selected radionuclides (gross alpha- and beta-particle activity [at 72 hours and 30 days], carbon-14, radium-226, radon-222, and uranium). Field measurements were made of selected physicochemical (relating to both physical and chemical) properties (oxidation-reduction potential, turbidity, dissolved-oxygen concentration, pH, specific conductance, water temperature, and alkalinity) and unfiltered sulfides. Similar to the results from the reconnaissance survey, physicochemical properties, major ions, and trace elements varied considerably. The ranges of selected physicochemical properties were as follows: oxidation-reduction potential ranged from -173 to 466 millivolts, dissolved oxygen ranged from less than 0.1 to 4.4 milligrams per liter, pH ranged from 7.2 to 7.8, specific conductance ranged from 439 to 724 microsiemens per centimeter at 25 degrees Celsius, and alkalinity ranged from 159 to 276 milligrams per liter as calcium carbonate. The largest ranges in concentration for filtered major ion constituents were obtained for cations sodium and calcium and for anions chloride and sulfate. Arsenic concentrations measured in samples from the 47 wells ranged from 1.6 to 23.5 micrograms per liter. The maximum concentration of arsenic (23.5 micrograms per liter) was measured in the source-water sample from well LJ-65-12-328. Quantifiable concentrations of barium, boron, lithium, molybdenum, and strontium were measured in all 47 filtered, source-water samples. Quantifiable concentrations of manganese were measured in 46 source-water samples, and an estimated concentration of manganese was measured in 1 sample. Chromium, iron, selenium, and vanadium were detected in 24 or more of the 47 source-water samples. Gross alpha-particle activities and beta-particle activities for all 47 samples were analyzed at 72 hours after sample collection and again at 30 days after sample collection, allowing for the measurement of the activity of short-lived isotopes. Gross alpha-particle activities reported in this report were not adjusted for activity contributions by radon or uranium and, therefore, are conservatively high estimates if compared to the U.S. Environmental Protection Agency National Primary Drinking Water Regulation for adjusted gross alpha-particle activity. The gross alpha-particle activities at 30 days in the samples ranged from R0.60 to 25.5 picocuries per liter and at 72 hours ranged from 2.58 to 39.7 picocuries per liter, and the "R" preceding the value of 0.60 picocuries per liter refers to a nondetected result less than the sample-specific critical level. Gross beta-particle activities measured at 30 days ranged from 1.17 to 14.4 picocuries per liter and at 72 hours ranged from 1.97 to 4.4 picocuries per liter. Filtered uranium was detected in quantifiable amounts in all of the 47 wells sampled. The uranium concentrations ranged from 0.03 to 42.7 micrograms per liter. One sample was analyzed for carbon-14, and the amount of modern atmospheric carbon was reported as 0.2 percent. Six source-water samples collected from municipal supply wells were analyzed for radium-226, and all of the concentrations were considered detectable concentrations (greater than their associated sample-specific critical level). Three source-water samples collected were analyzed for radon-222, and all of the concentrations were substantially greater than the associated sample-specific critical level.

Texas

Emerging prion disease drives host selection in a wildlife population

Infectious diseases are increasingly recognized as an important force driving population dynamics, conservation biology, and natural selection in wildlife populations. Infectious agents have been implicated in the decline of small or endangered populations and may act to constrain population size, distribution, growth rates, or migration patterns. Further, diseases may provide selective pressures that shape the genetic diversity of populations or species. Thus, understanding disease dynamics and selective pressures from pathogens is crucial to understanding population processes, managing wildlife diseases, and conserving biological diversity. There is ample evidence that variation in the prion protein gene (PRNP) impacts host susceptibility to prion diseases. Still, little is known about how genetic differences might influence natural selection within wildlife populations. Here we link genetic variation with differential susceptibility of white-tailed deer to chronic wasting disease (CWD), with implications for fitness and disease-driven genetic selection. We developed a single nucleotide polymorphism (SNP) assay to efficiently genotype deer at the locus of interest (in the 96th codon of the PRNP gene). Then, using a Bayesian modeling approach, we found that the more susceptible genotype had over four times greater risk of CWD infection; and, once infected, deer with the resistant genotype survived 49% longer (8.25 more months). We used these epidemiological parameters in a multi-stage population matrix model to evaluate relative fitness based on genotype-specific population growth rates. The differences in disease infection and mortality rates allowed genetically resistant deer to achieve higher population growth and obtain a long-term fitness advantage, which translated into a selection coefficient of over 1% favoring the CWD-resistant genotype. This selective pressure suggests that the resistant allele could become dominant in the population within an evolutionarily short time frame. Our work provides a rare example of a quantifiable disease-driven selection process in a wildlife population, demonstrating the potential for infectious diseases to alter host populations. This will have direct bearing on the epidemiology, dynamics, and future trends in CWD transmission and spread. Understanding genotype-specific epidemiology will improve predictive models and inform management strategies for CWD-affected cervid populations.

Ecological Applications

Hybridization decreases native cutthroat trout reproductive fitness

Examining natural selection in wild populations is challenging, but crucial to understanding many ecological and evolutionary processes. Additionally, in hybridizing populations, natural selection may be an important determinant of the eventual outcome of hybridization. We characterized several components of relative fitness in hybridizing populations of Yellowstone cutthroat trout and rainbow trout in an effort to better understand the prolonged persistence of both parental species despite predictions of extirpation. Thousands of genomic loci enabled precise quantification of hybrid status in adult and subsequent juvenile generations; a subset of those data also identified parent–offspring relationships. We used linear models and simulations to assess the effects of ancestry on reproductive output and mate choice decisions. We found a relatively low number of late-stage (F3+) hybrids and an excess of F2 juveniles relative to the adult generation in one location, which suggests the presence of hybrid breakdown decreasing the fitness of F2+ hybrids later in life. Assessments of reproductive output showed that Yellowstone cutthroat trout are more likely to successfully reproduce and produce slightly more offspring than their rainbow trout and hybrid counterparts. Mate choice appeared to be largely random, though we did find statistical support for slight female preference for males of similar ancestry. Together, these results show that native Yellowstone cutthroat trout are able to outperform rainbow trout in terms of reproduction and suggest that management action to exclude rainbow trout from spawning locations may bolster the now-rare Yellowstone cutthroat trout.

Molecular Ecology

SNP discovery in candidate adaptive genes using exon capture in a free-ranging alpine ungulate

Identification of genes underlying genomic signatures of natural selection is key to understanding adaptation to local conditions. We used targeted resequencing to identify SNP markers in 5321 candidate adaptive genes associated with known immunological, metabolic and growth functions in ovids and other ungulates. We selectively targeted 8161 exons in protein-coding and nearby 5′ and 3′ untranslated regions of chosen candidate genes. Targeted sequences were taken from bighorn sheep ( Ovis canadensis ) exon capture data and directly from the domestic sheep genome ( Ovis aries v. 3; oviAri3). The bighorn sheep sequences used in the Dall's sheep ( Ovis dalli dalli ) exon capture aligned to 2350 genes on the oviAri3 genome with an average of 2 exons each. We developed a microfluidic qPCR-based SNP chip to genotype 476 Dall's sheep from locations across their range and test for patterns of selection. Using multiple corroborating approaches ( lositan and bayescan ), we detected 28 SNP loci potentially under selection. We additionally identified candidate loci significantly associated with latitude, longitude, precipitation and temperature, suggesting local environmental adaptation. The three methods demonstrated consistent support for natural selection on nine genes with immune and disease-regulating functions (e.g. Ovar-DRA, APC, BATF2, MAGEB18), cell regulation signalling pathways (e.g. KRIT1, PI3K, ORRC3), and respiratory health (CYSLTR1). Characterizing adaptive allele distributions from novel genetic techniques will facilitate investigation of the influence of environmental variation on local adaptation of a northern alpine ungulate throughout its range. This research demonstrated the utility of exon capture for gene-targeted SNP discovery and subsequent SNP chip genotyping using low-quality samples in a nonmodel species.

Molecular Ecology Resources

How characteristic is the species characteristic selection scale?

Aim The importance of framing investigations of organism–environment relationships to interpret patterns at relevant spatial scales is increasingly recognized. However, most research related to environmental relationships is single-scaled, implicitly or explicitly assuming that a “species characteristic selection scale” exists. We tested the premise that a single characteristic scale exists to understand species–environment relationships within species by asking (a) what are the characteristic scales of species’ relationships with environmental predictors, and (b) is within-species, cross-predictor consistency in characteristic scales a general phenomenon. Location Nebraska, USA. Time period 2016. Major taxa studied Birds. Methods We used data from 86 species at > 500 locations to build hierarchical N-mixture models relating species abundance to land cover variables. By incorporating Bayesian latent indicator scale selection, we identified the spatial scales that best explain species–environment relationships with each land cover predictor. We quantified the extent of cross-predictor consistency in characteristic scales, and contrasted this to the expectation given a single species’ characteristic scale. Results We found no evidence for a characteristic spatial scale explaining all abundance–environment relationships within species, rather we found substantial variation in scale-dependence across multiple environmental attributes. Furthermore, 33% of species displayed evidence of multiple important spatial scales within environmental attributes. Major conclusions Within species there is little evidence for a single characteristic scale of environmental relationships and considerable variation in species’ scale dependencies. Because species may respond to multiple environmental attributes at different spatial scales, or single environmental attributes at multiple scales, we caution against any unoptimized single-scale studies. Our results demonstrate that until a framework is developed to predict the scales at which species respond to environmental characteristics, multi-scale investigations must be performed to identify and account for multi-scale dependencies. Natural selection acting on species’ response to distinct environmental attributes, rather than natural selection acting on species’ perception of spatial scales per se, may have shaped patterns of scale dependency and is an area ripe for investigation.

Nebraska

Model-based estimation of individual fitness

Fitness is the currency of natural selection, a measure of the propagation rate of genotypes into future generations. Its various definitions have the common feature that they are functions of survival and fertility rates. At the individual level, the operative level for natural selection, these rates must be understood as latent features, genetically determined propensities existing at birth. This conception of rates requires that individual fitness be defined and estimated by consideration of the individual in a modelled relation to a group of similar individuals; the only alternative is to consider a sample of size one, unless a clone of identical individuals is available. We present hierarchical models describing individual heterogeneity in survival and fertility rates and allowing for associations between these rates at the individual level. We apply these models to an analysis of life histories of Kittiwakes ( Rissa tridactyla ) observed at several colonies on the Brittany coast of France. We compare Bayesian estimation of the population distribution of individual fitness with estimation based on treating individual life histories in isolation, as samples of size one (e.g. McGraw & Caswell, 1996).

Brittany Coast