Colleen M. Handel

Extreme endurance flights by landbirds crossing the Pacific Ocean: ecological corridor rather than barrier?

Mountain ranges, deserts, ice fields and oceans generally act as barriers to the movement of land-dependent animals, often profoundly shaping migration routes. We used satellite telemetry to track the southward flights of bar-tailed godwits (Limosa lapponica baueri), shorebirds whose breeding and non-breeding areas are separated by the vast central Pacific Ocean. Seven females with surgically implanted transmitters flew non-stop 8117–11 680 km (10 153±1043 s.d.) directly across the Pacific Ocean; two males with external transmitters flew non-stop along the same corridor for 7008–7390 km. Flight duration ranged from 6.0 to 9.4 days (7.8±1.3 s.d.) for birds with implants and 5.0 to 6.6 days for birds with externally attached transmitters. These extraordinary non-stop flights establish new extremes for avian flight performance, have profound implications for understanding the physiological capabilities of vertebrates and how birds navigate, and challenge current physiological paradigms on topics such as sleep, dehydration and phenotypic flexibility. Predicted changes in climatic systems may affect survival rates if weather conditions at their departure hub or along the migration corridor should change. We propose that this transoceanic route may function as an ecological corridor rather than a barrier, providing a wind-assisted passage relatively free of pathogens and predators.

Hemispheric-scale wind selection facilitates bar-tailed godwit circum-migration of the Pacific

The annual 29 000 km long migration of the bar-tailed godwit, Limosa lapponica baueri, around the Pacific Ocean traverses what is arguably the most complex and seasonally structured atmospheric setting on Earth. Faced with marked variation in wind regimes and storm conditions across oceanic migration corridors, individuals must make critical decisions about when and where to fly during nonstop flights of a weeks duration or longer. At a minimum, their decisions will affect wind profitability and thus reduce energetic costs of migration; in the extreme, poor decisions or unpredictable weather events will risk survival. We used satellite telemetry to track the annual migration of 24 bar-tailed godwits and analysed their flight performance relative to wind conditions during three major migration legs between nonbreeding grounds in New Zealand and breeding grounds in Alaska. Because flight altitudes of birds en route were unknown, we modelled flight efficiency at six geopotential heights across each migratory segment. Birds selected departure dates when atmospheric conditions conferred the greatest wind assistance both at departure and throughout their flights. This behaviour suggests that there exists a cognitive mechanism, heretofore unknown among migratory birds, that allows godwits to assess changes in weather conditions that are linked (i.e. teleconnected) across widely separated atmospheric regions. Godwits also showed adaptive flexibility in their response not only to cues related to seasonal changes in macrometeorology, such as spatial shifting of storm tracks and temporal periods of cyclogenesis, but also to cues associated with stochastic events, especially at departure sites. Godwits showed limits to their response behaviours, however, especially relative to rapidly developing stochastic events while en route. We found that flight efficiency depended significantly upon altitude and hypothesize that godwits exhibit further adaptive flexibility by varying flight altitude en route to optimize flight efficiency. The Association for the Study of Animal Behaviour. Published by Elsevier Ltd.

A hierarchical model combining distance sampling and time removal to estimate detection probability during avian point counts

ABSTRACT Imperfect detection during animal surveys biases estimates of abundance and can lead to improper conclusions regarding distribution and population trends. Farnsworth et al. (2005) developed a combined distance-sampling and time-removal model for point-transect surveys that addresses both availability (the probability that an animal is available for detection; e.g., that a bird sings) and perceptibility (the probability that an observer detects an animal, given that it is available for detection). We developed a hierarchical extension of the combined model that provides an integrated analysis framework for a collection of survey points at which both distance from the observer and time of initial detection are recorded. Implemented in a Bayesian framework, this extension facilitates evaluating covariates on abundance and detection probability, incorporating excess zero counts (i.e. zero-inflation), accounting for spatial autocorrelation, and estimating population density. Species-specific characteristics, such as behavioral displays and territorial dispersion, may lead to different patterns of availability and perceptibility, which may, in turn, influence the performance of such hierarchical models. Therefore, we first test our proposed model using simulated data under different scenarios of availability and perceptibility. We then illustrate its performance with empirical point-transect data for a songbird that consistently produces loud, frequent, primarily auditory signals, the Golden-crowned Sparrow (Zonotrichia atricapilla); and for 2 ptarmigan species (Lagopus spp.) that produce more intermittent, subtle, and primarily visual cues. Data were collected by multiple observers along point transects across a broad landscape in southwest Alaska, so we evaluated point-level covariates on perceptibility (observer and habitat), availability (date within season and time of day), and abundance (habitat, elevation, and slope), and included a nested point-within-transect and park-level effect. Our results suggest that this model can provide insight into the detection process during avian surveys and reduce bias in estimates of relative abundance but is best applied to surveys of species with greater availability (e.g., breeding songbirds).

Epizootic of BEak DEformitiEs among WilD BirDs in alaska: an EmErging DisEasE in north amErica?

ABSTRACT. The sudden appearance of a large cluster of animals with gross abnormalities may signal a significant change in an ecosystem. We describe an unusual concentration of beak deformities that appear to have arisen rapidly within Alaska and now extend southward along the Pacific Coast. In Alaska we have documented 2,160 Black-capped Chickadees (Poecile atricapillus) and 435 individuals of 29 other species of birds, primarily during the past decade, with grossly overgrown and often crossed beaks. The annual prevalence of beak abnormalities among adult Black-capped Chickadees in south-central Alaska varied from 3.6% to 9.7% and averaged 6.5 ± 0.5% between 1999 and 2008. Only 0.05 ± 0.05% of nestlings and 0.3 ± 0.2% of juveniles <6 months old had abnormal beaks, which suggests that this is either a latent developmental or an acquired condition. We documented 80 cases in which a Black-capped Chickadee captured with an apparently normal beak was subsequently recaptured with a beak abnormality and 8 cases in which a beak deformity was no longer detectable upon recapture. Necropsy and histopathology of a sample of affected individuals provided no conclusive evidence of the etiology of this condition. Deformities appear to affect primarily the keratin layer of the beak and may result from abnormally rapid growth of the rhamphotheca. Some affected birds also exhibited lesions in other keratinized tissues of the skin, legs, feet, claws, and feathers, which may represent a systemic disorder or secondary conditions. Additional studies are currently underway to determine diagnostic signs and the underlying cause of this avian keratin disorder.

Beak Deformities in Northwestern Crows: Evidence of a Multispecies Epizootic

ABSTRACT. Beak abnormalities are rare among adult birds and, typically, are not widespread in a given population, within a region, or across multiple species. A high concentration of beak deformities was recently documented in Black-capped Chickadees (Poecile atricapillus) and other resident avian species in Alaska. We describe a parallel condition in Northwestern Crows (Corvus caurinus) that signals the emergence of a multispecies epizootic. On the basis of 186 Northwestern Crows captured at six sites in Alaska during 2007 and 2008, we estimated the prevalence of beak deformities in adults to be 16.9 ± 5.3%, the highest rate of gross deformities ever recorded in a wild bird population. Prevalence varied among sites and was as high as 36% on the Kenai Peninsula, which suggests possible epizootic clusters. We also documented beak abnormalities in an additional 148 Northwestern Crows in south-central and southeastern Alaska and in 64 crows near Vancouver, British Columbia, and Puget Sound, Washington, a region where both Northwestern Crows and American Crows (C. brachyrhynchos) occur. The increase in frequency and distribution of crows observed with abnormal beaks throughout the Pacific Northwest since the late 1990s indicates a geographic expansion of this problem. Affected crows exhibited elongated and often crossed beaks that were morphologically similar to deformities documented in Black-capped Chickadees and other species in Alaska over approximately the same period. Additional research is needed to determine the etiology and potential adverse effects on bird populations affected by this disorder.

Reviving common standards in point-count surveys for broad inference across studies

ABSTRACT We revisit the common standards recommended by Ralph et al. (1993, 1995a) for conducting point-count surveys to assess the relative abundance of landbirds breeding in North America. The standards originated from discussions among ornithologists in 1991 and were developed so that point-count survey data could be broadly compared and jointly analyzed by national data centers with the goals of monitoring populations and managing habitat. Twenty years later, we revisit these standards because (1) they have not been universally followed and (2) new methods allow estimation of absolute abundance from point counts, but these methods generally require data beyond the original standards to account for imperfect detection. Lack of standardization and the complications it introduces for analysis become apparent from aggregated data. For example, only 3% of 196,000 point counts conducted during the period 1992–2011 across Alaska and Canada followed the standards recommended for the count period and count radius. Ten-minute, unlimited-count-radius surveys increased the number of birds detected by >300% over 3-minute, 50-m-radius surveys. This effect size, which could be eliminated by standardized sampling, was ≥10 times the published effect sizes of observers, time of day, and date of the surveys. We suggest that the recommendations by Ralph et al. (1995a) continue to form the common standards when conducting point counts. This protocol is inexpensive and easy to follow but still allows the surveys to be adjusted for detection probabilities. Investigators might optionally collect additional information so that they can analyze their data with more flexible forms of removal and time-of-detection models, distance sampling, multiple-observer methods, repeated counts, or combinations of these methods. Maintaining the common standards as a base protocol, even as these study-specific modifications are added, will maximize the value of point-count data, allowing compilation and analysis by regional and national data centers.

THE RELATIVE IMPORTANCE OF NESTING AND FORAGING SITES IN SELECTION OF BREEDING TERRITORIES BY TOWNSEND'S WARBLERS

We investigated habitat selection by the Townsends Warbler (Dendroica town- sendi), a Neotropical-Nearctic migrant that breeds primarily in mature coniferous forests. From 1993 to 1994, we compared the features of habitat selected for nest sites and foraging sites with those selected for territories in mature, mixed coniferous-deciduous forests in south-central Alaska. We also tested the prediction that large conifers are selected for nesting and foraging sites. Females placed nests in relatively large white spruce (Picea glauca) and consistently chose nest trees that were within areas of higher densities of large white spruce than were generally available. Nesting areas also had a higher density of small white spruce than was generally available, which may have been important in concealing nests from pred- ators. Vegetation features selected for foraging differed from those selected for nesting and varied seasonally. Foraging was concentrated in medium-sized white spruce during the pre- hatching stage and became more generalized across coniferous and deciduous vegetation during the posthatching stage. Adults foraged in large white spruce in direct proportion to their availability on the study areas. Territories, which encompassed both nesting and for- aging areas, were heterogeneous in vegetation structure and floristics. Areas selected for territories clearly reflected availability of the large white spruce selected for nest sites but did not reflect selection of medium white spruce for foraging. Characteristics of habitats varied with specific resource needs, but the distribution of Townsends Warblers was most strongly related to specific habitat requirements for nest sites. Received 29 July 1996, accepted 5 May 1997.

Nesting ecology of Townsend's warblers in relation to habitat characteristics in a mature boreal forest

We investigated the nesting ecology of Townsends Warblers (Dendroica townsendi) from 1993-1995 in an unfragmented boreal forest along the lower slopes of the Chugach Mountains in southcentral Alaska. We examined habitat characteristics of nest sites in relation to factors influencing reproductive success. Almost all territory-holding males (98%, n = 40) were successful in acquiring mates. Nest success was 54% (n = 24 nests), with nest survivorship greater during incubation (87%) than during the nestling period (62%). Most nesting failure (80%) was attributable to predation, which occurred primarily during the nestling period. Fifty-five percent of nests containing nestlings were infested with the larvae of bird blow-flies (Protocalliphora braueri and P. spenceri), obligatory blood-feeding parasites. The combined effects of Protocalliphora infestation and inclement weather apparently resulted in nestling mortality in 4 of the 24 nests. Nests that escaped predation were placed in white spruce with larger diameter than those lost to predation; nests that escaped blow-fly parasitism were located higher in nest trees and in areas with lower densities of woody shrubs than those that were infested. The availability of potential nest sites with these key features may be important in determining reproductive success in Townsends Warblers.

Melanin-Based Color of Plumage: Role of Condition and of Feathers’ Microstructure

Whether melanin-based colors honestly signal a birds condition during the growth of feathers is controversial, and it is unclear if, or how, the physiological processes underlying melanogenesis or the role of the microstructure of feathers in imparting structural color to feathers may be adversely affected by condition. Here, we report results from two experiments designed to measure the effect of condition on expression of eumelanic and pheomelanic coloration in black-capped chickadees (Poecile atricapillus) and zebra finches (Taeniopygia guttata), respectively. In chickadees, we compared feathers of birds affected and unaffected by avian keratin disorder, whereas in zebra finches we compared feathers of controls with feathers of those subjected to an unpredictable food supply during development. In both cases, we found that control birds had brighter feathers (higher total reflectance) and more barbules, but similar densities of melanosomes. In addition, the microstructure of the feathers explained variation in color more strongly than did melanosome density. Together, these results suggest that melanin-based coloration may in part be condition-dependent, but that this may be driven by changes in keratin and feather development, rather than melanogenesis itself. Researchers should be cautious when assigning variation in melanin-based color to melanin alone and microstructure of the feather should be taken into account.

Wildlife health in a rapidly changing North: focus on avian disease

Climate‐related environmental changes have increasingly been linked to emerging infectious diseases in wildlife. The Arctic is facing a major ecological transition that is expected to substantially affect animal and human health. Changes in phenology or environmental conditions that result from climate warming may promote novel species assemblages as host and pathogen ranges expand to previously unoccupied areas. Recent evidence from the Arctic and subarctic suggests an increase in the spread and prevalence of some wildlife diseases, but baseline data necessary to detect and verify such changes are still lacking. Wild birds are undergoing rapid shifts in distribution and have been implicated in the spread of wildlife and zoonotic diseases. Here, we review evidence of current and projected changes in the abundance and distribution of avian diseases and outline strategies for future research. We discuss relevant climatic and environmental factors, emerging host–pathogen contact zones, the relationship between host condition and immune function, and potential wildlife and human health outcomes in northern regions.