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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).

Book chapter

Spatial partitioning and asymmetric hybridization among sympatric coastal steelhead trout (Oncorhynchus mykiss irideus), coastal cutthroat trout (O. clarki clarki) and interspecific hybrids

Hybridization between sympatric species provides unique opportunities to examine the contrast between mechanisms that promote hybridization and maintain species integrity. We surveyed hybridization between sympatric coastal steelhead (Oncorhynchus mykiss irideus) and coastal cutthroat trout (O. clarki clarki) from two streams in Washington State, Olsen Creek (256 individuals sampled) and Jansen Creek (431 individuals sampled), over a 3-year period. We applied 11 O. mykiss-specific nuclear markers, 11 O. c. clarki-specific nuclear markers and a mitochondrial DNA marker to assess spatial partitioning among species and hybrids and determine the directionality of hybridization. F1 and post-F1 hybrids, respectively, composed an average of 1.2% and 33.6% of the population sampled in Jansen Creek, and 5.9% and 30.4% of the population sampled in Olsen Creek. A modest level of habitat partitioning among species and hybrids was detected. Mitochondrial DNA analysis indicated that all F 1 hybrids (15 from Olsen Creek and five from Jansen Creek) arose from matings between steelhead females and cutthroat males implicating a sneak spawning behaviour by cutthroat males. First-generation cutthroat backcrosses contained O. c. clarki mtDNA more often than expected suggesting natural selection against F1 hybrids. More hybrids were backcrossed toward cutthroat than steelhead and our results indicate recurrent hybridization within these creeks. Age analysis demonstrated that hybrids were between 1 and 4 years old. These results suggest that within sympatric salmonid hybrid zones, exogenous processes (environmentally dependent factors) help to maintain the distinction between parental types through reduced fitness of hybrids within parental environments while divergent natural selection promotes parental types through distinct adaptive advantages of parental phenotypes.

Washington

Colonizing the world in spite of reduced MHC variation

Reduced immune gene diversity is thought to negatively affect the capacity of organisms to adapt to pathogen challenges, which represent a major force in natural selection. Genes of the Major Histocompatibility Complex (MHC) are the most widely invoked adaptive loci in conservation biology, and have become the most popular genetic markers to investigate pathogen-host interactions in vertebrates. Although MHC genes are the most polymorphic genes described in the vertebrate genome, the extent to which MHC diversity determines the long-term persistence of populations is, unclear and often debated, as recent studies have documented the occurrence of natural populations thriving even after a depletion of MHC diversity caused by genetic drift. Here, we show that some phylogenetically related species belonging to the Falco genus (Aves: Falconidae) present a dramatically low MHC variability that has not precluded, nevertheless, the successful colonization of almost all existing regions and habitats worldwide. We found evidence for two remarkably different patterns of MHC variation within the genus. While kestrels show a high MHC variation according to the general theory, falcons exhibit an ancestrally low intra- and inter-specific MHC allelic diversity. We provide compelling evidence that this pattern is not caused by the degeneration of functional genes into pseudogenes, the inadvertent analyses of paralogous MHC genes, or the devastating action of genetic drift. Instead, our results strongly support the idea of an evolutionary transition driven and maintained by natural selection from primarily highly variable towards low polymorphic, but functional and expressed, MHC genes with species-specific pathogen-recognition capabilities.

Journal of Evolutionary Biology

Genomic patterns of introgression in rainbow and westslope cutthroat trout illuminated by overlapping paired-end RAD sequencing

Rapid and inexpensive methods for genomewide single nucleotide polymorphism (SNP) discovery and genotyping are urgently needed for population management and conservation. In hybridized populations, genomic techniques that can identify and genotype thousands of species-diagnostic markers would allow precise estimates of population- and individual-level admixture as well as identification of 'super invasive' alleles, which show elevated rates of introgression above the genomewide background (likely due to natural selection). Techniques like restriction-site-associated DNA (RAD) sequencing can discover and genotype large numbers of SNPs, but they have been limited by the length of continuous sequence data they produce with Illumina short-read sequencing. We present a novel approach, overlapping paired-end RAD sequencing, to generate RAD contigs of >300–400 bp. These contigs provide sufficient flanking sequence for design of high-throughput SNP genotyping arrays and strict filtering to identify duplicate paralogous loci. We applied this approach in five populations of native westslope cutthroat trout that previously showed varying (low) levels of admixture from introduced rainbow trout (RBT). We produced 77 141 RAD contigs and used these data to filter and genotype 3180 previously identified species-diagnostic SNP loci. Our population-level and individual-level estimates of admixture were generally consistent with previous microsatellite-based estimates from the same individuals. However, we observed slightly lower admixture estimates from genomewide markers, which might result from natural selection against certain genome regions, different genomic locations for microsatellites vs. RAD-derived SNPs and/or sampling error from the small number of microsatellite loci (n = 7). We also identified candidate adaptive super invasive alleles from RBT that had excessively high admixture proportions in hybridized cutthroat trout populations.

Molecular Ecology

Genetic basis of differences in myxospore count between whirling disease-resistant and -susceptible strains of rainbow trout

We used a quantitative genetics approach and estimated broad sense heritability (h 2 b ) of myxospore count and the number of genes involved in myxospore formation to gain a better understanding of how resistance to Myxobolus cerebralis, the parasite responsible for whirling disease, is inherited in rainbow trout Oncorhynchus mykiss. An M. cerebralis-resistant strain of rainbow trout, the German Rainbow (GR), and a wild, susceptible strain of rainbow trout, the Colorado River Rainbow (CRR), were spawned to create 3 intermediate crossed populations (an F1 cross, F2 intercross, and a B2 backcross between the F1 and the CRR). Within each strain or cross, h 2 b was estimated from the between-family variance of myxospore counts using full-sibling families. Estimates of h 2 b and average myxospore counts were lowest in the GR strain, F1 cross, and F2 intercross (h 2 b = 0.34, 0.42, and 0.34; myxospores fish −1 = 275, 9566, and 45780, respectively), and highest in the B2 backcross and CRR strain (h 2 b = 0.93 and 0.89; myxospores fish −1 = 97865 and 187595, respectively). Comparison of means and a joint-scaling test suggest that resistance alleles arising from the GR strain are dominant to susceptible alleles from the CRR strain. Resistance was retained in the intermediate crosses but decreased as filial generation number increased (F2) or backcrossing occurred (B2). The estimated number of segregating loci responsible for differences in myxospore count in the parental strains was 9 ± 5. Our results indicate that resistance to M. cerebralis is a heritable trait within these populations and would respond to either artificial selection in hatcheries or natural selection in the wild.

Diseases of Aquatic Organisms

Diurnal habitat selection and survival of elk neonates

Natural selection should favor development of behaviors that increase survival, including juvenile survival. Habitat characteristics (e.g., hiding cover, forage quality and availability), maternal habitat selection, and microhabitat selection by the calf may influence elk ( Cervus canadensis ) calf survival and recruitment. We assessed diurnal microhabitat selection of bed sites by elk calves and calf-rearing areas selected by adult females to determine if these characteristics were associated with calf survival. We radio-tagged 33 elk calves in west-central New Mexico in 2011 and 55 calves in north-central New Mexico in 2012. We tracked calves daily to locate calf bedding sites and collected data on selected physical features and vegetation characteristics at used and paired random sites. The paired differences in these characteristics were then associated with calf fate. At the calf selection scale, for every 1-m difference in the distance to nearest concealment cover, the odds that a site was from a surviving calf increased by 7.8%. Habitat selection by adult females also was associated with calf survival. The odds of a bed site being from a surviving calf increased by 1.9% for every 1% difference in percentage of grass cover. High levels of concealment cover at the bed site were related to calf survival status. When we expanded selection to the adult level, females with surviving calves selected areas with higher grass cover, suggesting adult selection for higher forage availability while still providing high concealment cover for the calf.

Southwestern Naturalist

Evidence of economical territory selection in a cooperative carnivore

As an outcome of natural selection, animals are probably adapted to select territories economically by maximizing benefits and minimizing costs of territory ownership. Theory and empirical precedent indicate that a primary benefit of many territories is exclusive access to food resources, and primary costs of defending and using space are associated with competition, travel and mortality risk. A recently developed mechanistic model for economical territory selection provided numerous empirically testable predictions. We tested these predictions using location data from grey wolves ( Canis lupus ) in Montana, USA. As predicted, territories were smaller in areas with greater densities of prey, competitors and low-use roads, and for groups of greater size. Territory size increased before decreasing curvilinearly with greater terrain ruggedness and harvest mortalities. Our study provides evidence for the economical selection of territories as a causal mechanism underlying ecological patterns observed in a cooperative carnivore. Results demonstrate how a wide range of environmental and social conditions will influence economical behaviour and resulting space use. We expect similar responses would be observed in numerous territorial species. A mechanistic approach enables understanding how and why animals select particular territories. This knowledge can be used to enhance conservation efforts and more successfully predict effects of conservation actions.

Montana

Equations for estimating selected streamflow statistics in Rhode Island

Regional regression equations were developed for estimating selected natural—unaffected by alteration—streamflows of specific flow durations and low-flow frequency statistics for ungaged stream sites in Rhode Island. Selected at-site streamflow statistics are provided for 41 long-term streamgages, 21 short-term streamgages, and 135 partial-record stations in Rhode Island, eastern Connecticut, and southeastern and south-central Massachusetts. The regression equations for estimating selected streamflow statistics and the at-site statistics estimated for each of the 197 sites may be used by Federal, State, and local water managers in addressing water issues in and near Rhode Island. Multiple and simple linear regression equations were developed to estimate the 99-, 98-, 95-, 90-, 85-, 80-, 75-, 70-, 60-, 50-, 40-, 30-, 25-, 20-, 15-, 10-, 5-, 2-, and 1-percent flow durations and the 7Q2 (7-day, 2-year) and 7Q10 (7-day, 10-year) low-flow-frequency statistics. An additional 49 selected statistics, for which regression equations were not developed, also were estimated for the long- and short-term streamgages and partial-record stations for flow durations between the 99.99 and 0.01 percent and for the mean annual, mean monthly, and median monthly streamflows. A total of 70 selected streamflow statistics were estimated for 41 long-term streamgages, 21 short-term streamgages, and 135 partial-record stations in and near Rhode Island. Estimates of the long-term streamflow statistics for the 21 short-term streamgages and 135 partial-record stations were developed by the Maintenance of Variance Extension, type 1 (MOVE.1), record-extension technique. The equations used to estimate selected streamflow statistics were developed by relating the 19 flow-duration and 2 low-flow-frequency statistics to 31 different basin characteristics (physical, land-cover, and climatic) at the 41 long-term and 19 of 21 short-term streamgages (a total of 60 streamgages) in and near Rhode Island. The 135 partial-record stations were not used in the regression analyses. The regression analyses were done by using a user-weighted least-squares technique in the weighted-multiple-linear regression program for the 90- to 1-percent flow-duration statistics. For the 99-, 98-, and 95-percent flow durations and the 7Q2 and 7Q10 statistics, left-censored regression analyses were used to account for zero flows at a few streamgages. The regression analyses determined that two basin characteristics—drainage area and stream density—were the only significant explanatory variables for 16 of the 19 flow-duration and the 2 low-flow regression equations. For the 10-, 15-, and 20-percent flow-duration regression equations, drainage area was the only significant explanatory variable. The standard error of the estimate for the 21 regression equations ranged from 17.58 to 141.83 percent. The 99- to 85-percent flow durations and the low-flow statistics 7Q2 and 7Q10 had the highest standard errors of the estimate, ranging from 48.68 to 141.83 percent. The standard error of the estimate for the medium- to high-flow statistics—the 80- to 1-percent flow durations—ranged from 17.58 to 37.65 percent, with the standard errors for the 60- to 1-percent flow durations all being less than about 21 percent. Data also are provided to allow the user to calculate the 90-percent prediction intervals for the 21 streamflow statistics. The equations, which are based on data from streams with little to no flow alterations, will provide an estimate of the natural flows for a selected site. They will not estimate flows for altered sites with dams, surface-water withdrawals, groundwater withdrawals (pumping wells), diversions, and wastewater discharges. If the equations are used to estimate streamflow statistics for altered sites, the user should adjust the flow estimates for the alterations. The regression equations should be used only for ungaged sites with drainage areas between 0.52 and 294 square miles and stream densities between 0.94 and 3.49 miles per square mile; these are the ranges of the explanatory variables in the equations.

Rhode Island

Linking physical monitoring to coho and Chinook salmon populations in the Redwood Creek Watershed, California—Summary of May 3–4, 2012 Workshop

On Thursday, May 3, 2012, a science workshop was held at the Redwood National and State Parks (RNSP) office in Arcata, California, with researchers and resource managers working in RNSP to share data and expert opinions concerning salmon populations and habitat in the Redwood Creek watershed. The focus of the workshop was to discuss how best to synthesize physical and biological data related to the freshwater and estuarine phases of salmon life cycles in order to increase the understanding of constraints on salmon populations. The workshop was hosted by the U.S. Geological Survey (USGS) Status and Trends (S&T) Program National Park Monitoring Project ( http://www.fort.usgs.gov/brdscience/ParkMonitoring.htm ), which supports USGS research on priority topics (themes) identified by the National Park Service (NPS) Inventory and Monitoring Program (I&M) and S&T. The NPS has organized more than 270 parks with significant natural resources into 32 Inventory and Monitoring (I&M) Networks ( http://science.nature.nps.gov/im/networks.cfm ) that share funding and core professional staff to monitor the status and long-term trends of selected natural resources ( http://science.nature.nps.gov/im/monitor ). All 32 networks have completed vital signs monitoring plans (available at http://science.nature.nps.gov/im/monitor/MonitoringPlans.cfm ), containing background information on the important resources of each park, conceptual models behind the selection of vital signs for monitoring the condition of natural resources, and the selection of high priority vital signs for monitoring. Vital signs are particular physical, chemical, and biological elements and processes of park ecosystems that represent the overall health or condition of the park, known or hypothesized effects of stressors, or elements that have important human values (Fancy and others, 2009). Beginning in 2009, the I&M program funded projects to analyze and synthesize the biotic and abiotic data generated by vital signs monitoring and previous in-park natural resource monitoring and inventories to provide useful information, models, and tools to park managers for addressing resource management issues. The workshop described in this report is an element of the project funded by USGS NPS-I&M program to conduct a synthesis of salmon-related datasets in the Klamath (KLMN) and San Francisco Bay Area (SFAN) networks of national parks. The synthesis focused on four park units: Redwood National Park (KLMN), Point Reyes National Seashore, Muir Woods National Monument, and Golden Gate National Recreation Area (SFAN).

California

Arsenic and radionuclide occurrence and relation to geochemistry in groundwater of the Gulf Coast Aquifer System in Houston, Texas, 2007–11

The U.S. Geological Survey (USGS), in cooperation with the City of Houston, began a study in 2007 to determine concentrations, spatial extent, and associated geochemical conditions that might be conducive for mobility and transport of selected naturally occurring trace elements and radionuclides in the Gulf Coast aquifer system in Houston, Texas. Water samples were collected from 91 municipal supply wells completed in the Evangeline and Chicot aquifers of the Gulf Coast aquifer system in northeastern, northwestern, and southwestern Houston; hereinafter referred to as northeast, northwest and southwest Houston areas. Wells were sampled in three phases: (1) 28 municipal supply wells were sampled during 2007–8, (2) 60 municipal supply wells during 2010, and (3) 3 municipal supply wells during December 2011. During each phase of sampling, samples were analyzed for major ions, selected trace elements, and radionuclides. At a subset of wells, concentrations of arsenic species and other radionuclides (carbon-14, radium-226, radium-228, radon-222, and tritium) also were analyzed. Selected physicochemical properties were measured in the field at the time each sample was collected, and oxidation-reduction potential and unfiltered sulfides also were measured at selected wells. The source-water (the raw, ambient water withdrawn from municipal supply wells prior to water treatment) samples were collected for assessment of aquifer conditions in order to provide community water-system operators information that could be important when they make decisions about which treatment processes to apply before distributing finished drinking water. Geochemical conditions of groundwater of the Gulf Coast aquifer system are suitable in some instances for release of arsenic and radionuclides from aquifer materials. Recent changes to the U.S. Environmental Protection Agency (EPA) primary drinking-water regulations for arsenic and a selected number of natural radionuclides have highlighted the necessity for municipal supply system managers to be aware of the occurrence and distribution of these constituents in their source water. Concentrations of arsenic ranged from 0.58 to 23.5 micrograms per liter (μg/L), with relatively low median and 75th percentile concentrations (2.7 and 3.6 μg/L, respectively). The gross alpha-particle activity completed within 72 hours after sample collection ranged from R-1.1 (nondetect where the result was below the sample specific critical level) to 39.7 picocuries per liter (pCi/L), with a median of 10.3 pCi/L. After 30 days, the gross alpha-particle activities in the 91 samples ranged from R-0.94 to 25.5 pCi/L, with a median of 5.60 pCi/L. Concentrations of uranium ranged from less than 0.02 to 42.7 μg/L, with a median value of 1.69 μg/L and a 75th-percentile value of 6.48 μg/L. The maximum concentrations of radium-226 and combined radium (sum of radium-226 plus radium-228) were 4.34 pCi/L and 3.23 pCi/L, respectively. Aquifer major-ion geochemistry was characterized and shown to contain three chemical types of water as grouped by a simplified predominant cation and anion classification system: (1) calcium- bicarbonate type, (2) sodium-bicarbonate type, and (3) sodium-chloride type. Aquifer geochemistry also was characterized into four reduction-oxidation (redox) categories: (1) oxic, (2) suboxic, (3) mixed, and (4) anoxic. Within the anoxic category, groundwater was further characterized into four presumed predominant reduction processes: (1) iron or sulfate or both [Fe(III)/SO 4 ] reducing, (2) iron [Fe(III)] reducing, (3) iron and sulfate [Fe(III)-SO 4 ] reducing, or (4) methanogenic, as defined by composition of redox species. The oxic category was associated with calcium-bicarbonate-type water, and the methanogenic-anoxic process was associated exclusively with the sodium-bicarbonate-type water. The species of arsenic and the dominant radionuclide present were associated with specific redox categories. Arsenate was associated primarily with oxic water and did not exceed 3.5 µg/L, whereas arsenite was associated with iron-reducing, anoxic water samples and, at the highest concentrations, occurred in sulfate-reducing, anoxic; methanogenic-anoxic; or both water samples. Uranium was associated exclusively with the oxic water, whereas the highest concentrations of combined radium were associated with the iron-reducing, anoxic water. The gross alpha-particle activity was greatest in the oxic waters where the source of the radioactivity was the uranium. Associated geochemical conditions conducive for mobility of arsenic and radionuclides and their spatial and vertical extent in the Gulf Coast aquifer system in Houston are important aspects to the areal management of the municipal groundwater supplies in Houston. Ongoing research is seeking to define chemical or geological factors that are the optimal indicators for elevated concentrations of these naturally occurring constituents.

Texas

Dispersal and selection mediate hybridization between a native and invasive species

Hybridization between native and non-native species has serious biological consequences, but our understanding of how dispersal and selection interact to influence invasive hybridization is limited. Here, we document the spread of genetic introgression between a native (Oncorhynchus clarkii) and invasive (Oncorhynchus mykiss) trout, and identify the mechanisms influencing genetic admixture. In two populations inhabiting contrasting environments, non-native admixture increased rapidly from 1984 to 2007 and was driven by surprisingly consistent processes. Individual admixture was related to two phenotypic traits associated with fitness: size at spawning and age of juvenile emigration. Fish with higher non-native admixture were larger and tended to emigrate at a younger age—relationships that are expected to confer fitness advantages to hybrid individuals. However, strong selection against non-native admixture was evident across streams and cohorts (mean selection coefficient against genotypes with non-native alleles (s) ¼ 0.60; s.e. ¼ 0.10). Nevertheless, hybridization was promoted in both streams by the continuous immigration of individuals with high levels of non-native admixture from other hybrid source populations. Thus, antagonistic relationships between dispersal and selection are mediating invasive hybridization between these fish, emphasizing that data on dispersal and natural selection are needed to fully understand the dynamics of introgression between native and non-native species. .

Proceedings of the Royal Society B

Vive la résistance: genome-wide selection against introduced alleles in invasive hybrid zones

Evolutionary and ecological consequences of hybridization between native and invasive species are notoriously complicated because patterns of selection acting on non-native alleles can vary throughout the genome and across environments. Rapid advances in genomics now make it feasible to assess locus-specific and genome-wide patterns of natural selection acting on invasive introgression within and among natural populations occupying diverse environments. We quantified genome-wide patterns of admixture across multiple independent hybrid zones of native westslope cutthroat trout and invasive rainbow trout, the world's most widely introduced fish, by genotyping 339 individuals from 21 populations using 9380 species-diagnostic loci. A significantly greater proportion of the genome appeared to be under selection favouring native cutthroat trout (rather than rainbow trout), and this pattern was pervasive across the genome (detected on most chromosomes). Furthermore, selection against invasive alleles was consistent across populations and environments, even in those where rainbow trout were predicted to have a selective advantage (warm environments). These data corroborate field studies showing that hybrids between these species have lower fitness than the native taxa, and show that these fitness differences are due to selection favouring many native genes distributed widely throughout the genome.

Proceedings of the Royal Society B: Biological Sci

The aerosphere as a network connector of organisms and their diseases

Aeroecological processes, especially powered flight of animals, can rapidly connect biological communities across the globe. This can have profound consequences for evolutionary diversification, energy and nutrient transfers, and the spread of infectious diseases. The latter is of particular consequence for human populations, since migratory birds are known to host diseases which have a history of transmission into domestic poultry or even jumping to human hosts. In this chapter, we present a scenario under which a highly pathogenic avian influenza (HPAI) strain enters North America from East Asia via post-molting waterfowl migration. We use an agent-based model (ABM) to simulate the movement and disease transmission among 10 6 generalized waterfowl agents originating from ten molting locations in eastern Siberia, with the HPAI seeded in only ~10 2 agents at one of these locations. Our ABM tracked the disease dynamics across a very large grid of sites as well as individual agents, allowing us to examine the spatiotemporal patterns of change in virulence of the HPAI infection as well as waterfowl host susceptibility to the disease. We concurrently simulated a 12-station disease monitoring network in the northwest USA and Canada in order to assess the potential efficacy of these sites to detect and confirm the arrival of HPAI. Our findings indicated that HPAI spread was initially facilitated but eventually subdued by the migration of host agents. Yet, during the 90-day simulation, selective pressures appeared to have distilled the HPAI strain to its most virulent form (i.e., through natural selection), which was counterbalanced by the host susceptibility being conversely reduced (i.e., through genetic predisposition and acquired immunity). The monitoring network demonstrated wide variation in the utility of sites; some were clearly better at providing early warnings of HPAI arrival, while sites further from the disease origin exposed the selective dynamics which slowed the spread of the disease albeit with the result of passing highly virulent strains into southern wintering locales (where human impacts are more likely). Though the ABM presented had generalized waterfowl migration and HPAI disease dynamics, this exercise demonstrates the power of such simulations to examine the extremely large and complex processes which comprise aeroecology. We offer insights into how such models could be further parameterized to represent HPAI transmission risks as well as how ABMs could be applied to other aeroecological questions pertaining to individual-based connectivity.

Book chapter

Moose habitat selection and fitness consequences during two critical winter tick life stages in Vermont, United States

The moose ( Alces alces ) is a charismatic species in decline across much of their southern distribution in North America. In the northeastern United States, much of the reduction has been attributed to winter tick ( Dermacentor albipictus ) infestations. Winter ticks are fairly immobile throughout all life stages, and therefore their distribution patterns at any given time are shaped largely by the occurrence of moose across the landscape during the peak of two critical time periods: fall questing (when ticks latch onto moose) and spring drop-off (when engorged female ticks detach from moose). We used recent land cover and lidar data within a dynamic occupancy modeling framework to estimate first-order habitat selection (use vs. non-use) of female moose ( n = 74) during the tick questing and drop-off periods. Patch extinction and colonization rates between the fall questing and spring drop-off periods were strongly influenced by habitat and elevation, but these effects were diminished during the fall questing period when moose were more active across the landscape. From the fall questing period to the spring drop-off period, patches where colonization was high and extinction was low had higher proportions of young (shrub/forage) mixed forest at higher elevations. Further, we evaluated the fitness consequences of habitat selection by adult females during the fall questing period, when females and their calves acquire ticks. We compared Resource Selection Functions (RSF) for five females that successfully reared a calf to age 1 with five females whose calves perished due to ticks. Adult female moose whose offspring perished selected habitats in the fall that spatially coincided with areas of high occupancy probability during the spring tick drop-off period. In contrast, adult female moose whose offspring survived selected areas where the probability of occupancy during the spring drop-off was low; at present, natural selection may favor female adults who do not select the same habitats in fall as in spring. Our model coefficients and mapped results define “hotspots” that are likely encouraging the deleterious effects of the tick-moose cycle. These findings fill knowledge gaps about moose habitat selection that may improve the effectiveness of management aimed at reversing declining population trends.

Vermont

The geometry of reaction norms yields insights on classical fitness functions for Great Lakes salmon

Life history theory examines how characteristics of organisms, such as age and size at maturity, may vary through natural selection as evolutionary responses that optimize fitness. Here we ask how predictions of age and size at maturity differ for the three classical fitness functions–intrinsic rate of natural increase r , net reproductive rate R 0 , and reproductive value V x −for semelparous species. We show that different choices of fitness functions can lead to very different predictions of species behavior. In one’s efforts to understand an organism’s behavior and to develop effective conservation and management policies, the choice of fitness function matters. The central ingredient of our approach is the maturation reaction norm (MRN), which describes how optimal age and size at maturation vary with growth rate or mortality rate. We develop a practical geometric construction of MRNs that allows us to include different growth functions (linear growth and nonlinear von Bertalanffy growth in length) and develop two-dimensional MRNs useful for quantifying growth-mortality trade-offs. We relate our approach to Beverton-Holt life history invariants and to the Stearns-Koella categorization of MRNs. We conclude with a detailed discussion of life history parameters for Great Lakes Chinook Salmon and demonstrate that age and size at maturity are consistent with predictions using R 0 (but not r or V x ) as the underlying fitness function.

Great Lakes

Atlantic salmon brood stock management and breeding handbook

Anadromus runs of Atlantic salmon have been restored to the Connecticut, Merrimack, Pawcatuck, Penobscot, and St. Croix rivers in New England by the stocking of more than 8 million smolts since 1948. Fish-breeding methods have been developed that minimize inbreeding and domestication and enhance natural selection. Methods are available to advance the maturation of brood stock, control the sex of production lots and store gametes. Current hatchery practices emphasize the use of sea-run brood stock trapped upon return to the rivers and a limited number of captive brood stock and rejuvenated kelts. Fish are allowed to mature naturally, after which they are spawned and incubated artificially. Generally, 1-year smolts are produced, and excess fish are stocked as fry in headwater streams. Smolts are stocked during periods of rising water in spring. Self-release pools are planned that enable smolts to choose the emigration time. Culturists keep good records that permit evaluation of the performance of strains and the effects of breeding practices. As Atlantic salmon populations expand, culturists must use sound breeding methods that enhance biotic potential while maintaining genetic diversity and protecting unique gene pools.

Biological Report

Bioavailable iron production in airborne mineral dust: Controls by chemical composition and solar flux

A large part of oceanic biological production is limited by the scarcity of dissolved iron. Mineral dust aerosol , processed under acidic atmospheric conditions , is the primary natural source of bioavailable iron to oceanic life. However, synergistic and antagonistic effects of non-Fe-containing minerals on atmospheric processing of Fe-containing minerals and Fe solubilization are poorly understood. The current study focuses on mineralogical influences of non-Fe-bearing semiconductor minerals, such as titanium dioxide (TiO 2 ), on the dissolution of iron in selected natural mineral dust aerosols under atmospherically relevant conditions. Further, the role of elevated Ti concentrations in dust is evaluated using magnetite , a proxy for Fe(II) containing minerals, under both dark and light conditions. Our results highlight that relatively higher Ti:Fe ratios, regardless of their total Fe content, enhances the total iron dissolution in mineral dust aerosols as well as in magnetite. Moreover, elevated Ti percentages also yield high Fe(II) fractions in mineral dust systems under dark conditions. Upon irradiation however, dissolved Fe(II) is suppressed by high Ti levels due to the involvement of photochemical redox cycling reactions with hydroxyl radicals ( • OH). These synergistic and antagonistic effects of Ti are further evaluated by altering the chemical composition of natural dusts with artificially added anatase (TiO 2 ) and synthetic amorphous titania . The current study reveals important mineralogical controls by non-Fe-bearing minerals on dust iron dissolution to better understand global iron mobilization.

Atmospheric Environment

The evolution of different maternal investment strategies in two closely related desert vertebrates

We compared egg size phenotypes and tested several predictions from the optimal egg size (OES) and bet-hedging theories in two North American desert-dwelling sister tortoise taxa, Gopherus agassizii and G. morafkai , that inhabit different climate spaces: relatively unpredictable and more predictable climate spaces, respectively. Observed patterns in both species differed from the predictions of OES in several ways. Mean egg size increased with maternal body size in both species. Mean egg size was inversely related to clutch order in G. agassizii , a strategy more consistent with the within-generation hypothesis arising out of bet-hedging theory or a constraint in egg investment due to resource availability, and contrary to theories of density dependence, which posit that increasing hatchling competition from later season clutches should drive selection for larger eggs. We provide empirical evidence that one species, G. agassizii , employs a bet-hedging strategy that is a combination of two different bet-hedging hypotheses. Additionally, we found some evidence for G. morafkai employing a conservative bet-hedging strategy. (e.g., lack of intra- and interclutch variation in egg size relative to body size). Our novel adaptive hypothesis suggests the possibility that natural selection favors smaller offspring in late-season clutches because they experience a more benign environment or less energetically challenging environmental conditions (i.e., winter) than early clutch progeny, that emerge under harsher and more energetically challenging environmental conditions (i.e., summer). We also discuss alternative hypotheses of sexually antagonistic selection, which arise from the trade-offs of son versus daughter production that might have different optima depending on clutch order and variation in temperature-dependent sex determination (TSD) among clutches. Resolution of these hypotheses will require long-term data on fitness of sons versus daughters as a function of incubation environment, data as yet unavailable for any species with TSD.

Arizona, California