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Improved arrival-date estimates of Arctic-breeding Dunlin ( Calidris alpina arcticola )
The use of stable isotopes in animal ecology depends on accurate descriptions of isotope dynamics within individuals. The prevailing assumption that laboratory-derived isotopic parameters apply to free-living animals is largely untested. We used stable carbon isotopes (δ 13 C) in whole blood from migratory Dunlin ( Calidris alpina arcticola ) to estimate an in situ turnover rate and individual diet-switch dates. Our in situ results indicated that turnover rates were higher in free-living birds, in comparison to the results of an experimental study on captive Dunlin and estimates derived from a theoretical allometric model. Diet-switch dates from all 3 methods were then used to estimate arrival dates to the Arctic; arrival dates calculated with the in situ turnover rate were later than those with the other turnover-rate estimates, substantially so in some cases. These later arrival dates matched dates when local snow conditions would have allowed Dunlin to settle, and agreed with anticipated arrival dates of Dunlin tracked with light-level geolocators. Our study presents a novel method for accurately estimating arrival dates for individuals of migratory species in which return dates are difficult to document. This may be particularly appropriate for species in which extrinsic tracking devices cannot easily be employed because of cost, body size, or behavioral constraints, and in habitats that do not allow individuals to be detected easily upon first arrival. Thus, this isotopic method offers an exciting alternative approach to better understand how species may be altering their arrival dates in response to changing climatic conditions.
Fast and efficient: postnatal growth and energy expenditure in an Arctic-breeding waterbird, the Red-throated Loon ( Gavia stellata )
Environmental conditions can exert a strong influence on the growth and energy demands of chicks. We hypothesized that postnatal growth in a cold, aquatic environment would require a high level of energy metabolism in semiprecocial Red-throated Loon ( Gavia stellata ) chicks. We measured body-mass growth and daily energy expenditure (DEE) of free-ranging chicks in the Arctic. We used daily gains in body mass and DEE to estimate daily metabolizable energy (DME, kJ day -1 ) and total metabolizable energy (TME, kJ chick -1 ). Chicks gained body mass quickly, with a logistic growth rate constant 57% greater than the allometric prediction, yet were at only 60% of adult body mass at fledging. Males grew at a rate similar to that of females but for a slightly longer duration and so reached an asymptotic body mass 23% greater, and tarsus length 8% longer, than that of females. Chick growth performance was similar between first- and second-hatched chicks within broods of 2, which suggests that food availability was not limited. DEE increased in proportion to body mass, and DME peaked at 1,214 kJ day -1 on day 25 posthatching. Over the average 49-day postnatal period, TME was 49.0 MJ, which is within the range of error of the allometric prediction. Parents provided 58.6 MJ as food to meet this energy requirement. Given this chick energy requirement and the range of energy content of prey observed in the chick diet, selecting prey with higher energy content would greatly reduce adult provisioning effort. Red-throated Loon chicks did not have a high postnatal energy requirement, but rather grew quickly and fledged at a small size-with the effect of reducing the length of the postnatal period and, consequently, parental energy investment in chicks.
Morphological divergence in a continental adaptive radiation: South American ovenbirds of the genus Cinclodes
Cinclodes is an ecologically diverse genus of South American passerine birds and represents a case of continental adaptive radiation along multiple axes. We investigated morphological diversification in Cinclodes using a comprehensive set of morphometric measurements of study skins. Principal component analysis identified 2 primary axes of morphological variation: one describing body size and a second capturing differences in wing-tip shape and toe length. Phylogenetic analyses of the first principal component suggest an early divergence of Cinclodes into 2 main clades characterized by large and small body sizes. We suggest that 2 morphological outliers within these main clades ( C. antarcticus and C. palliatus ) may be cases of island gigantism and that a third ( C. patagonicus ) may reflect ecological character displacement. Despite its ecological and physiological diversity, the genus Cinclodes does not appear to show morphological diversity beyond what is typical of other avian genera.
Redefining reproductive success in songbirds: Moving beyond the nest success paradigm
One of the most commonly estimated parameters in studies of songbird ecology is reproductive success, as a measure of either individual fitness or population productivity. Traditionally, the “success” in reproductive success refers to whether, or how many, nestlings leave nests. Here, we advocate that “reproductive success” in songbirds be redefined as full-season productivity, or the number of young raised to independence from adult care in a breeding season. A growing body of evidence demonstrates interdependence between nest success and fledgling survival, and emphasizes that data from either life stage alone can produce misleading measures of individual fitness and population productivity. Nest success, therefore, is an insufficient measure of reproductive success, and songbird ecology needs to progress beyond this long-standing paradigm. Full-season productivity, an evolutionarily rational measure of reproductive success, provides the framework for appropriately addressing unresolved questions about the adaptive significance of many breeding behaviors and within which effective breeding-grounds conservation and management can be designed.
Life-history tradeoffs and reproductive cycles in Spotted Owls
The study of tradeoffs among life-history traits has long been key to understanding the evolution of life-history strategies. However, more recently, evolutionary ecologists have realized that reproductive costs have the potential to influence population dynamics. Here, we tested for costs of reproduction in the California Spotted Owl ( Strix occidentalis occidentalis ), and assessed whether costs of reproduction in year t − 1 on reproduction in year t could be responsible for regionally synchronized biennial cycles in reproductive output. Logistic regression analysis and multistate mark–recapture models with state uncertainty revealed that breeding reduced the likelihood of reproducing in the subsequent year by 16% to 38%, but had no influence on subsequent survival. We also found that costs of reproduction in year t − 1 were correlated with climatic conditions in year t , with evidence of higher costs during the dry phase of the El Niño–Southern Oscillation. Using a simulation-based population model, we showed that strong reproductive costs had the potential to create biennial cycles in population-level reproductive output; however, estimated costs of reproduction appeared to be too small to explain patterns observed in Spotted Owls. In the absence of strong reproductive costs, we hypothesize that observed natural cycles in the reproductive output of Spotted Owls are related to as-yet-unmeasured, regionally concordant fluctuations in environmental conditions or prey resources. Despite theoretical evidence for demographic effects, our analyses illustrate that linking tradeoffs to actual changes in population processes will be challenging because of the potential confounding effects of individual and environmental variation.
In-flight turbulence benefits soaring birds
Birds use atmospheric updrafts to subsidize soaring flight. We observed highly variable soaring flight by Black Vultures ( Coragyps atratus ) and Turkey Vultures ( Cathartes aura ) in Virginia, USA, that was inconsistent with published descriptions of terrestrial avian flight. Birds engaging in this behavior regularly deviated vertically and horizontally from linear flight paths. We observed the soaring flight behavior of these 2 species to understand why they soar in this manner and when this behavior occurs. Vultures used this type of soaring mainly at low altitudes (<50 m), along forest edges, and when conditions were poor for thermal development. Because of the tortuous nature of this flight, we describe it as “contorted soaring.” The primary air movement suitable to subsidize flight at this altitude and under these atmospheric conditions is small-scale, shear-induced turbulence, which our results suggest can be an important resource for soaring birds because it permits continuous subsidized flight when other types of updraft are not available.
Dispersal and survival of a polygynandrous passerine
Although sex biases in survival and dispersal are thought to be linked to avian mating systems, little is known about these demographic patterns in less common mating strategies such as polygynandry. We investigated breeding-site fidelity, natal philopatry, and apparent survival of the polygynandrous Smith's Longspur ( Calcarius pictus ) over a 7-yr period at 2 areas in Alaska's Brooks Range. We used capture–recapture histories of 243 color-banded adults and 431 juveniles to estimate annual survival and determined dispersal patterns from 34 adults that were found breeding within the study areas over multiple years. Most adults (88%) returned to nest in the same breeding neighborhood as in previous years; mean dispersal distance was 300.9 ± 74.2 m and did not differ between sexes. Juveniles exhibited low natal philopatry; only 4% of banded hatch-year birds were resighted as adults during subsequent years. Those that did return dispersed, on average, 1,674.4 ± 465.8 m from their natal nests ( n = 6). Model-averaged survival estimates indicated that annual survival of adult females (50–58%) was only slightly lower than that of males (60–63%); juvenile survival was 41% but was paired with a low (13%) encounter probability. We attribute the lack of sex bias in adult dispersal to this species' polygynandrous mating strategy. Within this system, there are multiple mates within a breeding neighborhood. We argue that natural selection may favor females that remain on the same, familiar breeding site, because they do not have to disperse to a new area to find a suitable mate. Dispersal among breeding populations most likely occurs by juveniles returning as adults. Our findings support hypotheses that suggest a relationship between dispersal and mating strategy and provide some of the first insight into the demographic patterns of a polygynandrous passerine.
Counterintuitive roles of experience and weather on migratory performance
Migration allows animals to live in resource-rich but seasonally variable environments. Because of the costs of migration, there is selective pressure to capitalize on variation in weather to optimize migratory performance. To test the degree to which migratory performance (defined as speed of migration) of Golden Eagles ( Aquila chrysaetos ) was determined by age- and season-specific responses to variation in weather, we analyzed 1,863 daily tracks ( n = 83 migrant eagles) and 8,047 hourly tracks ( n = 83) based on 15 min GPS telemetry data from Golden Eagles and 277 hourly tracks based on 30 s data ( n = 37). Spring migrant eagles traveled 139.75 ± 82.19 km day −1 (mean ± SE; n = 57) and 25.59 ± 11.75 km hr −1 ( n = 55). Autumn migrant eagles traveled 99.14 ± 59.98 km day −1 ( n = 26) and 22.18 ± 9.18 km hr −1 ( n = 28). Weather during migration varied by season and by age class. During spring, best-supported daily and hourly models of 15 min data suggested that migratory performance was influenced most strongly by downward solar radiation and that older birds benefited less from flow assistance (tailwinds). During autumn, best-supported daily and hourly models of 15 min data suggested that migratory performance was influenced most strongly by south–north winds and by flow assistance, again less strongly for older birds. In contrast, models for hourly performance based on data collected at 30 s intervals were not well described by a single model, likely reflecting eagles' rapid responses to the many weather conditions they experienced. Although daily speed of travel was similar for all age classes, younger birds traveled at faster hourly speeds than did adults. Our analyses uncovered strong, sometimes counterintuitive, relationships among weather, experience, and migratory flight, and they illustrate the significance of factors other than age in determining migratory performance.
Logistic quantile regression provides improved estimates for bounded avian counts: A case study of California Spotted Owl fledgling production
Counts of avian fledglings, nestlings, or clutch size that are bounded below by zero and above by some small integer form a discrete random variable distribution that is not approximated well by conventional parametric count distributions such as the Poisson or negative binomial. We developed a logistic quantile regression model to provide estimates of the empirical conditional distribution of a bounded discrete random variable. The logistic quantile regression model requires that counts are randomly jittered to a continuous random variable, logit transformed to bound them between specified lower and upper values, then estimated in conventional linear quantile regression, repeating the 3 steps and averaging estimates. Back-transformation to the original discrete scale relies on the fact that quantiles are equivariant to monotonic transformations. We demonstrate this statistical procedure by modeling 20 years of California Spotted Owl fledgling production (0−3 per territory) on the Lassen National Forest, California, USA, as related to climate, demographic, and landscape habitat characteristics at territories. Spotted Owl fledgling counts increased nonlinearly with decreasing precipitation in the early nesting period, in the winter prior to nesting, and in the prior growing season; with increasing minimum temperatures in the early nesting period; with adult compared to subadult parents; when there was no fledgling production in the prior year; and when percentage of the landscape surrounding nesting sites (202 ha) with trees ≥25 m height increased. Changes in production were primarily driven by changes in the proportion of territories with 2 or 3 fledglings. Average variances of the discrete cumulative distributions of the estimated fledgling counts indicated that temporal changes in climate and parent age class explained 18% of the annual variance in owl fledgling production, which was 34% of the total variance. Prior fledgling production explained as much of the variance in the fledgling counts as climate, parent age class, and landscape habitat predictors. Our logistic quantile regression model can be used for any discrete response variables with fixed upper and lower bounds.
Niche shifts and energetic condition of songbirds in response to phenology of food-resource availability in a high-elevation sagebrush ecosystem
Seasonal fluctuations in food availability can affect diets of consumers, which in turn may influence the physiological state of individuals and shape intra- and inter-specific patterns of resource use. High-elevation ecosystems often exhibit a pronounced seasonal “pulse” in productivity, although few studies document how resource use and energetic condition by avian consumers change in relation to food-resource availability in these ecosystems. We tested the hypothesis that seasonal increases (pulses) in food resources in high-elevation sagebrush ecosystems result in 2 changes after the pulse, relative to the before-pulse period: (1) reduced diet breadth of, and overlap between, 2 sympatric sparrow species; and (2) enhanced energetic condition in both species. We tracked breeding-season diets using stable isotopes and energetic condition using plasma metabolites of Brewer's Sparrows ( Spizella breweri ), Vesper Sparrows ( Pooecetes gramineus ), and their food resources during 2011, and of only Brewer's Sparrows and their food resources during 2013. We quantify diet breadth and overlap between both species, along with coincident physiological consequences of temporal changes in resource use. After invertebrate biomass increased following periods of rainfall in 2011, dietary breadth decreased by 35% in Brewer's Sparrows and by 48% in Vesper Sparrows, while dietary overlap decreased by 88%. Energetic condition of both species increased when dietary overlap was lower and diet breadth decreased, after the rapid rise of food-resource availability. However, energetic condition of Brewer's Sparrows remained constant in 2013, a year with low precipitation and lack of a strong pulse in food resources, even though the species' dietary breadth again decreased that year. Our results indicate that diet breadth and overlap in these sparrow species inhabiting sagebrush ecosystems generally varied as predicted in relation to intra- and interannual changes in food resources, and this difference in diet was associated with improved energetic condition of sparrows at least in one year.
A field ornithologist’s guide to genomics: Practical considerations for ecology and conservation
Vast improvements in sequencing technology have made it practical to simultaneously sequence millions of nucleotides distributed across the genome, opening the door for genomic studies in virtually any species. Ornithological research stands to benefit in three substantial ways. First, genomic methods enhance our ability to parse and simultaneously analyze both neutral and non-neutral genomic regions, thus providing insight into adaptive evolution and divergence. Second, the sheer quantity of sequence data generated by current sequencing platforms allows increased precision and resolution in analyses. Third, high-throughput sequencing can benefit applications that focus on a small number of loci that are otherwise prohibitively expensive, time-consuming, and technically difficult using traditional sequencing methods. These advances have improved our ability to understand evolutionary processes like speciation and local adaptation, but they also offer many practical applications in the fields of population ecology, migration tracking, conservation planning, diet analyses, and disease ecology. This review provides a guide for field ornithologists interested in incorporating genomic approaches into their research program, with an emphasis on techniques related to ecology and conservation. We present a general overview of contemporary genomic approaches and methods, as well as important considerations when selecting a genomic technique. We also discuss research questions that are likely to benefit from utilizing high-throughput sequencing instruments, highlighting select examples from recent avian studies.
Fifty-seventh supplement to the American Ornithologists' Union Check-list of North American Birds
This is the 16th supplement since publication of the 7th edition of the Check-list of North American Birds (American Ornithologists' Union [AOU] 1998). It summarizes decisions made between April 15, 2015, and April 15, 2016, by the AOU's Committee on Classification and Nomenclature—North and Middle America. The Committee has continued to operate in the manner outlined in the 42nd Supplement (AOU 2000).
Environmental and ecological conditions at Arctic breeding sites have limited effects on true survival rates of adult shorebirds
Many Arctic shorebird populations are declining, and quantifying adult survival and the effects of anthropogenic factors is a crucial step toward a better understanding of population dynamics. We used a recently developed, spatially explicit Cormack–Jolly–Seber model in a Bayesian framework to obtain broad-scale estimates of true annual survival rates for 6 species of shorebirds at 9 breeding sites across the North American Arctic in 2010–2014. We tested for effects of environmental and ecological variables, study site, nest fate, and sex on annual survival rates of each species in the spatially explicit framework, which allowed us to distinguish between effects of variables on site fidelity versus true survival. Our spatially explicit analysis produced estimates of true survival rates that were substantially higher than previously published estimates of apparent survival for most species, ranging from S = 0.72 to 0.98 across 5 species. However, survival was lower for the arcticola subspecies of Dunlin ( Calidris alpina arcticola ; S = 0.54), our only study taxon that migrates through the East Asian–Australasian Flyway. Like other species that use that flyway, arcticola Dunlin could be experiencing unsustainably low survival rates as a result of loss of migratory stopover habitat. Survival rates of our study species were not affected by timing of snowmelt or summer temperature, and only 2 species showed minor variation among study sites. Furthermore, although previous reproductive success, predator abundance, and the availability of alternative prey each affected survival of one species, no factors broadly affected survival across species. Overall, our findings of few effects of environmental or ecological variables suggest that annual survival rates of adult shorebirds are generally robust to conditions at Arctic breeding sites. Instead, conditions at migratory stopovers or overwintering sites might be driving adult survival rates and should be the focus of future studies.
Combining multiple sources of data to inform conservation of Lesser Prairie-Chicken populations
Conservation of small populations is often based on limited data from spatially and temporally restricted studies, resulting in management actions based on an incomplete assessment of the population drivers. If fluctuations in abundance are related to changes in weather, proper management is especially important, because extreme weather events could disproportionately affect population abundance. Conservation assessments, especially for vulnerable populations, are aided by a knowledge of how extreme events influence population status and trends. Although important for conservation efforts, data may be limited for small or vulnerable populations. Integrated population models maximize information from various sources of data to yield population estimates that fully incorporate uncertainty from multiple data sources while allowing for the explicit incorporation of environmental covariates of interest. Our goal was to assess the relative influence of population drivers for the Lesser Prairie-Chicken ( Tympanuchus pallidicinctus ) in the core of its range, western and southern Kansas, USA. We used data from roadside lek count surveys, nest monitoring surveys, and survival data from telemetry monitoring combined with climate (Palmer drought severity index) data in an integrated population model. Our results indicate that variability in population growth rate was most influenced by variability in juvenile survival. The Palmer drought severity index had no measurable direct effects on adult survival or mean number of offspring per female; however, there were declines in population growth rate following severe drought. Because declines in population growth rate occurred at a broad spatial scale, declines in response to drought were likely due to decreases in chick and juvenile survival rather than emigration outside of the study area. Overall, our model highlights the importance of accounting for environmental and demographic sources of variability, and provides a thorough method for simultaneously evaluating population demography in response to long-term climate effects.
Adaptive influence of extrinsic and intrinsic factors on variation of incubation periods among tropical and temperate passerines
Understanding intrinsic (physiological) and extrinsic (e.g., temperature) causes of variation in embryonic development time (incubation period) is important because they can have different impacts on individual quality. Robert Ricklefs and colleagues have argued that longer incubation periods result primarily from intrinsic physiological programs that increase individual quality and adult survival. They claim that incubation periods are largely invariant and that extrinsic factors like temperature have little impact. We have argued that adult survival may be a cause rather than a consequence of much of the variation in embryonic development time. A reduction in extrinsic sources of annual adult mortality (e.g., migration, predation, nonbreeding-season mortality) favors reduced parental effort during incubation to minimize costs to future reproduction and survival. Reduced parental effort, in turn, manifests as cooler average egg temperatures that yield longer incubation periods. Ricklefs and colleagues mischaracterized our hypothesis and deconstructed their own incorrect version, while also making some incorrect statements. We show that reevaluation of previous evidence provided by this group actually supports a role of egg temperature for the variation in incubation periods. We also summarize other observational and experimental evidence that incubation periods are not invariant and that egg temperature has a strong causal influence on variation within and among species. In fact, egg temperature explains ∼60% of the difference in incubation periods among species. The remaining ∼40% reflects intrinsic physiological programs and other factors, potentially providing intrinsic benefits. Ultimately, annual adult mortality explains substantial variation in parental effort and egg temperature, and the latter strongly explains variation in incubation periods. Both intrinsic programs and extrinsic temperature effects need to be considered in attempts to understand incubation strategies.
The relative importance of intrinsic and extrinsic drivers to population growth vary among local populations of Greater Sage-Grouse: An integrated population modeling approach
Consideration of ecological scale is fundamental to understanding and managing avian population growth and decline. Empirically driven models for population dynamics and demographic processes across multiple spatial scales can be powerful tools to help guide conservation actions. Integrated population models (IPMs) provide a framework for better parameter estimation by unifying multiple sources of data (e.g., count and demographic data). Hierarchical structure within such models that include random effects allow for varying degrees of data sharing across different spatiotemporal scales. We developed an IPM to investigate Greater Sage-Grouse ( Centrocercus urophasianus ) on the border of California and Nevada, known as the Bi-State Distinct Population Segment. Our analysis integrated 13 years of lek count data ( n > 2,000) and intensive telemetry (VHF and GPS; n > 350 individuals) data across 6 subpopulations. Specifically, we identified the most parsimonious models among varying random effects and density-dependent terms for each population vital rate (e.g., nest survival). Using a joint likelihood process, we integrated the lek count data with the demographic models to estimate apparent abundance and refine vital rate parameter estimates. To investigate effects of climatic conditions, we extended the model to fit a precipitation covariate for instantaneous rate of change ( r ). At a metapopulation extent (i.e. Bi-State), annual population rate of change λ ( e r ) did not favor an overall increasing or decreasing trend through the time series. However, annual changes in λ were driven by changes in precipitation (one-year lag effect). At subpopulation extents, we identified substantial variation in λ and demographic rates. One subpopulation clearly decoupled from the trend at the metapopulation extent and exhibited relatively high risk of extinction as a result of low egg fertility. These findings can inform localized, targeted management actions for specific areas, and status of the species for the larger Bi-State.
Biological responses of Crested and Least auklets to volcanic destruction of nesting habitat in the Aleutian Islands, Alaska
Crested Auklets ( Aethia cristatella ) and Least Auklets ( A. pusilla ) are crevice-nesting birds that breed in large mixed colonies at relatively few sites in the Aleutian Island archipelago, Bering Sea, Gulf of Alaska, and Sea of Okhotsk. Many of these colonies are located on active volcanic islands. The eruption of Kasatochi volcano, in the central Aleutians, on August 7, 2008, completely buried all crevice-nesting seabird habitat on the island. This provided an opportunity to examine the response of a large, mixed auklet colony to a major geological disturbance. Time-lapse imagery of nesting habitat indicated that both species returned to the largest pre-eruption colony site for several years, but subsequently abandoned it within 5 yr after the eruption. In 2010, a rockfall site in a cove north of the old colony site began to accumulate talus, and groups of auklets were observed using the site in 2011. Use of the new colony appeared to coincide with the abandonment of the old colony site by both species, though surface counts suggested that Least Auklets shifted to the new colony sooner than Crested Auklets. At-sea surveys of seabirds before and after the eruption indicated that both Crested and Least auklets shifted their at-sea distributions from the waters around Kasatochi Island to nearby Koniuji Island. In combination, at-sea counts and colony time-lapse imagery indicated that Crested and Least auklets using Kasatochi responded to the volcanic disturbance and complete loss of nesting habitat at the main colony on Kasatochi with dispersal either to newly created habitat on Kasatochi or to an alternate colony on a nearby island.