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Landbird migration in the American West: Recent progress and future research directions

Our knowledge of avian behaviors during the nonbreeding period still lags behind that of the breeding season, but the last decade has witnessed a proliferation in research that has yielded significant progress in understanding migration patterns of North American birds. And, although historically the great majority of migration research has been conducted in the eastern half of the continent, there has been much recent progress on aspects of avian migration in the West. In particular, expanded use of techniques such as radar, plasma metabolites, mist-netting, count surveys, stable isotopes, genetic data, and animal tracking, coupled with an increase in multi-investigator collaborations, have all contributed to this growth of knowledge. There is increasing recognition that migration is likely the most limiting time of year for migratory birds, increasing the importance of continuing to decipher patterns of stopover ecology, identifying critical stopover habitats, and documenting migration routes in the diverse and changing landscapes of the American West. Here, we review and briefly synthesize the latest findings and advances in avian migration and consider research needs to guide future research on migration in the West. ?? 2009 by The Cooper Ornithological Society. All rights reserved.

The Condor↗

Environmental and human influences on trumpeter swan habitat occupancy in Alaska

Approximately 70-80% of the entire population of the Trumpeter Swan (Cygnus huccinator) depends for reproduction on wetlands in Alaska. This makes the identification of important habitat features and the effects of human interactions important for the species' long-term management. We analyzed the swan's habitat preferences in five areas throughout the state and found that swan broods occupied some wetland types, especially larger closed-basin wetlands such as lakes and ponds, at rates much higher than they occupied other wetland types, such as shrubby or forested wetlands. We also found a negative effect of transportation infrastructure on occupancy by broods in and around the Minto Flats State Game Refuge, Kenai National Wildlife Refuge, and Tetlin National Wildlife Refuge. This finding is of particular interest because much of the Minto Flats refuge has recently been licensed for oil and gas exploration and parts of the Kenai refuge have been developed in the past. We also investigated the potential effects of the shrinkage of closed-basin ponds on habitat occupancy by nesting Trumpeter Swans. We compared nesting swans' use of ponds with changes in the ponds' size and other characteristics from 1982 to 1996 and found no relationships between occupancy and changes in pond size. However, we believe that the recent and rapid growth of Trumpeter Swan populations in Alaska may become limited by available breeding habitat, and anthropogenic and climate-induced changes to the swan's breeding habitats have the potential to limit future production. ?? 2009 by The Cooper Ornithological Society. All rights reserved.

Condor↗

Avian response to wildfire in interior Columbia basin shrubsteppe

Wildfire and conversion of sagebrush (Artemisia spp.) shrublands to cheatgrass (Bromus tectorum) grasslands is a serious threat to the shrubsteppe ecosystem, but few studies have documented wildfire's effects on birds with multiple years of pre- and post-fire data. Using data from avian point counts recorded 4 years before and 7 years after a large-scale, severe wildfire in the Columbia Basin of south-central Washington, we found significant effects of fire on population trends or mean abundance of nearly all species investigated. The Sage Sparrow (Amphispiza belli), a sagebrush obligate, was decreasing at a high rate both pre- and post-fire. Among species inhabiting more open shrubsteppe or grasslands, the mean abundance of three (Grasshopper Sparrow, Ammodramus savannarum; Western Meadowlark, Sturnella neglecta; Vesper Sparrow, Pooecetes gramineus) was lower post-fire and one (Lark Sparrow, Chondestes grammacus) showed an initial, but short-lived, increase post-fire before dropping below pre-fire levels. Only one (Horned Lark, Eremophila alpestris) increased steadily post-fire and had higher post-fire mean abundance. ?? 2009 by The Cooper Ornithological Society. All rights reserved.

Condor↗

Probability of detection of nests and implications for survey design

Surveys based on double sampling include a correction for the probability of detection by assuming complete enumeration of birds in an intensively surveyed subsample of plots. To evaluate this assumption, we calculated the probability of detecting active shorebird nests by using information from observers who searched the same plots independently. Our results demonstrate that this probability varies substantially by species and stage of the nesting cycle but less by site or density of nests. Among the species we studied, the estimated single-visit probability of nest detection during the incubation period varied from 0.21 for the White-rumped Sandpiper (Calidris fuscicollis), the most difficult species to detect, to 0.64 for the Western Sandpiper (Calidris mauri), the most easily detected species, with a mean across species of 0.46. We used these detection probabilities to predict the fraction of persistent nests found over repeated nest searches. For a species with the mean value for detectability, the detection rate exceeded 0.85 after four visits. This level of nest detection was exceeded in only three visits for the Western Sandpiper, but six to nine visits were required for the White-rumped Sandpiper, depending on the type of survey employed. Our results suggest that the double-sampling method's requirement of nearly complete counts of birds in the intensively surveyed plots is likely to be met for birds with nests that survive over several visits of nest searching. Individuals with nests that fail quickly or individuals that do not breed can be detected with high probability only if territorial behavior is used to identify likely nesting pairs. ?? The Cooper Ornithological Society, 2009.

Condor↗

Exposure of nonbreeding migratory shorebirds to cholinesterase-inhibiting contaminants in the western hemisphere

Migratory shorebirds frequently forage and roost in agricultural habitats, where they may be exposed to cholinesterase-inhibiting pesticides. Exposure to organophosphorus and carbamate compounds, common anti-cholinesterases, can cause sublethal effects, even death. To evaluate exposure of migratory shorebirds to organophosphorus and carbamates, we sampled birds stopping over during migration in North America and wintering in South America. We compared plasma cholinesterase activities and body masses of individuals captured at sites with no known sources of organophosphorus or carbamates to those captured in agricultural areas where agrochemicals were recommended for control of crop pests. In South America, plasma acetylcholinesterase and butyrylcholinesterase activity in Buff-breasted Sandpipers was lower at agricultural sites than at reference sites, indicating exposure to organophosphorus and carbamates. Results of plasma cholinesterase reactivation assays and foot-wash analyses were inconclusive. A meta-analysis of six species revealed no widespread effect of agricultural chemicals on cholinesterase activity. however, four of six species were negative for acetylcholinesterase and one of six for butyrylcholinesterase, indicating negative effects of pesticides on cholinesterase activity in a subset of shorebirds. Exposure to cholinesterase inhibitors can decrease body mass, but comparisons between treatments and hemispheres suggest that agrochemicals did not affect migratory shorebirds' body mass. Our study, one of the first to estimate of shorebirds' exposure to cholinesterase-inhibiting pesticides, suggests that shorebirds are being exposed to cholinesterase- inhibiting pesticides at specific sites in the winter range but not at migratory stopover sites. future research should examine potential behavioral effects of exposure and identify other potential sitesand levels of exposure. ?? The Cooper Ornithological Society 2010.

Condor↗

Space use and habitat selection of migrant and resident American Avocets in San Francisco Bay

San Francisco Bay is a wintering area for shorebirds, including American Avocets (Recurvirostra americana). Recently, a new resident population of avocets has emerged, presumably because of the development of tidal marshes into salt-evaporation ponds. In habitat restoration now underway, as many as 90% of salt ponds will be restored to tidal marsh. However, it is unknown if wintering and resident avocets coexist and if their requirements for space and habitat differ, necessitating different management for their populations to be maintained during restoration. We captured and radio-marked wintering avocets at a salt pond and a tidal flat to determine their population status (migrant or resident) and examine their space use and habitat selection. Of the radio-marked avocets, 79% were migrants and 21% were residents. At the salt pond, residents' fidelity to their location of capture was higher, and residents moved less than did migrants from the same site. Conversely, on the tidal flat, fidelity of residents to their site of capture was lower, and residents' home ranges were larger than those of migrants from the same site. Habitat selection of migrants and residents differed little; however, capture site influenced habitat selection far more than the birds' status as migrants or residents. Our study suggests that individual avocets have high site fidelity while wintering in San Francisco Bay, although the avocet as a species is plastic in its space use and habitat selection. This plasticity may allow wintering migrant and resident avocets to adapt to habitat change in San Francisco Bay. ?? The Cooper Ornithological Society 2010.

Condor↗

Are there optimal densities for prairie birds?

The major forces of food and predation shape fitness-enhancing decisions of birds at all stages of their life cycles. During the breeding season, birds can minimize nest loss due to predation by selecting sites with a lower probability of predation. To understand the environmental and social aspects and consequences of breedingsite selection in prairie birds, we explored variation in nest-survival patterns of the Lark Bunting (Calamospiza melanocorys) in the shortgrass prairie region of North America. Over four breeding seasons, we documented the survival of 405 nests, conducted 60 surveys to estimate bird densities, and measured several vegetative features to describe habitat structure in 24 randomly selected study plots. Nest survival varied with the buntings' density as described by a quadratic polynomial, increasing with density below 1.5 birds ha -1 and decreasing with density between 1.5 and 3 birds ha -1 , suggesting that an optimal range of densities favors reproductive success of the Lark Bunting, which nests semi-colonially. Nest survival also increased with increasing vegetation structure of study plots and varied with age of the nest, increasing during early incubation and late in the nestling stage and declining slightly from mid-incubation to the middle of the nestling period. The existence of an optimal range of densities in this semi-colonial species can be elucidated by the "commodity-selection hypothesis" at low densities and density dependence at high densities. ?? The Cooper Ornithological Society 2010.

Condor↗

Trace elements have limited utility for studying migratory connectivity in shorebirds that winter in Argentina

Trace-element analysis has been suggested as a tool for the study of migratory connectivity because (1) trace-element abundance varies spatially in the environment, (2) trace elements are assimilated into animals' tissues through the diet, and (3) current technology permits the analysis of multiple trace elements in a small tissue sample, allowing the simultaneous exploration of several elements. We explored the potential of trace elements (B, Na, Mg, Al, Si, P, S, K, Ca, Ti, Cr, Mn, Ni, Cu, Zn, As, Sr, Cs, Hg, Tl, Pb, Bi, Th, and U) to clarify the migratory connectivity of shorebirds that breed in North America and winter in southern South America. We collected 66 recently replaced secondary feathers from Red Knots (Calidris canutus) at three sites in Patagonia and 76 from White-rumped Sandpipers (C. fuscicollis) at nine sites across Argentina. There were significant differences in trace-element abundance in shorebird feathers grown at different nonbreeding sites, and annual variability within a site was small compared to variability among sites. Across Argentina, there was no large-scale gradient in trace elements. The lack of such a gradient restricts the application of this technique to questions concerning the origin of shorebirds to a small number of discrete sites. Furthermore, our results including three additional species, the Pectoral Sandpiper (C. melanotos), Wilson's Phalarope (Phalaropus tricolor), and Collared Plover (Charadrius collaris), suggest that trace-element profiles change as feathers age. Temporal instability of trace-element values could undermine their application to the study of migratory connectivity in shorebirds. ?? The Cooper Ornithological Society 2010.

Condor↗

Seasonal movements, winter range use, and migratory connectivity of the Black Oystercatcher

The Black Oystercatcher (Haematopus bachmani) is an intertidal obligate along North America's Pacific coast and a species of high conservation concern (population size 8900-11 000 individuals). Understanding birds' movements and space use throughout the annual cycle has become paramount in the face of changing environmental conditions, and intertidal species may be particularly vulnerable to habitat change due to anticipated sea-level rise associated with climate change and increasing coastal development. Conservation of the Black Oystercatcher is hindered by a lack of information on the species' nonbreeding distribution, seasonal movements, and habitat connectivity. Using satellite (n = 19) and VHF (n = 19) radio transmitters, we tracked Black Oystercatchers from five breeding sites (Vancouver Island, British Columbia; Kodiak Island, Prince William Sound, Middleton Island, and Juneau, Alaska) through one and one half annual cycles (May 2007-Dec 2008). We documented medium- to long-distance migration (range of migration distance 130-1667 km) in three populations (Prince William Sound, Middleton Island, and Juneau) and year-round residency in two others (Kodiak and Vancouver Island). We observed variation in the timing and length of migration by study site, and individual birds demonstrated fidelity to breeding and nonbreeding sites. We did not observe strong migratory connectivity. Migratory oystercatchers distributed themselves widely along the coasts of British Columbia and southeast Alaska during winter. Results provide baseline information on the Black Oystercatcher's movements and space use throughout the annual cycle. ?? 2010 The Cooper Ornithological Society.

Condor↗

Parental investment decisions in response to ambient nest-predation risk versus actual predation on the prior nest

Theory predicts that parents should invest less in dependent offspring with lower reproductive value, such as those with a high risk of predation. Moreover, high predation risk can favor reduced parental activity when such activity attracts nest predators. Yet, the ability of parents to assess ambient nest-predation risk and respond adaptively remains unclear, especially where nest-predator assemblages are diverse and potentially difficult to assess. We tested whether variation in parental investment by a multi-brooded songbird (Brewer's Sparrow, Spizella breweri) in an environment (sagebrush steppe) with diverse predators was predicted by ambient nest-predation risk or direct experience with nest predation. Variation among eight sites in ambient nest-predation risk, assayed by daily probabilities of nest predation, was largely uncorrelated across four years. In this system risk may therefore be unpredictable, and aspects of parental investment (clutch size, egg mass, incubation rhythms, nestling-feeding rates) were not related to ambient risk. Moreover, investment at first nests that were successful did not differ from that at nests that were depredated, suggesting parents could not assess and respond to territorylevel nest-predation risk. However, parents whose nests were depredated reduced clutch sizes and activity at nests attempted later in the season by increasing the length of incubation shifts (on-bouts) and recesses (off-bouts) and decreasing trips to feed nestlings. In this unpredictable environment parent birds may therefore lack sufficient cues of ambient risk on which to base their investment decisions and instead rely on direct experience with nest predation to inform at least some of their decisions. ?? 2010 The Cooper Ornithological Society.

Condor↗

Latitudinal variation in reproductive strategies by the migratory Louisiana Waterthrush

We evaluated hypotheses that seek to explain breeding strategies of the Louisiana Waterthrush (Parkesia motacilla) that vary across a latitudinal gradient. On the basis of data from 418 nests of color-banded individuals in southwestern Pennsylvania and 700 km south in the Georgia Piedmont, we found that clutch size in replacement nests and probability of renesting were significantly greater in Pennsylvania (clutch size 4.4; renesting probability 0.66) than in Georgia (clutch size 3.8; renesting probability 0.54). Contrasts of the remaining measures of breeding were not statistically significant, and, in particular, mean daily nest survival in the two study areas was nearly identical (0.974 in Pennsylvania; 0.975 in Georgia). An individual-based model of fecundity (i.e., number of fledged young per adult female), predicted that approximately half of the females in both Pennsylvania and Georgia fledge at least one young, and mean values for fecundity in Pennsylvania and Georgia were 2.28 and 1.91, respectively. On the basis of greater support for the food-limitation hypothesis than for the season-length hypothesis, the trade-off between breeding in a region with more food but making a longer migration may be greater for waterthrushes breeding farther north than for those breeding farther south. ?? The Cooper Ornithological Society 2011.

Condor↗

Intercontinental gene flow among western arctic populations of Lesser Snow Geese

Quantifying the spatial genetic structure of highly vagile species of birds is important in predicting their degree of population demographic and genetic independence during changing environmental conditions, and in assessing their abundance and distribution. In the western Arctic, Lesser Snow Geese ( Chen caerulescens caerulescens ) provide an example useful for evaluating spatial population genetic structure and the relative contribution of male and female philopatry to breeding and wintering locales. We analyzed biparentally inherited microsatellite loci and maternally inherited mtDNA sequences from geese breeding at Wrangel Island (Russia) and Banks Island (Canada) to estimate gene flow among populations whose geographic overlap during breeding and winter differ. Significant differences in the frequencies of mtDNA haplotypes contrast with the homogeneity of allele frequencies for microsatellite loci. Coalescence simulations revealed high variability and asymmetry between males and females in rates and direction of gene flow between populations. Our results highlight the importance of wintering areas to demographic independence and spatial genetic structure of these populations. Male-mediated gene flow among the populations on northern Wrangel Island, southern Wrangel Island, and Banks Island has been substantial. A high rate of female-mediated gene flow from southern Wrangel Island to Banks Island suggests that population exchange can be achieved when populations winter in a common area. Conversely, when birds from different breeding populations do not share a common wintering area, the probability of population exchange is likely to be dramatically reduced.

The Condor↗

The influence of fine-scale habitat features on regional variation in population performance of alpine White-tailed Ptarmigan

It is often assumed (explicitly or implicitly) that animals select habitat features to maximize fitness. However, there is often a mismatch between preferred habitats and indices of individual and population measures of performance. We examined the influence of fine-scale habitat selection on the overall population performance of the White-tailed Ptarmigan (Lagopus leucura), an alpine specialist, in two subdivided populations whose habitat patches are configured differently. The central region of Vancouver Island, Canada, has more continuous and larger habitat patches than the southern region. In 2003 and 2004, using paired logistic regression between used (n = 176) and available (n = 324) sites, we identified food availability, distance to standing water, and predator cover as preferred habitat components . We then quantified variation in population performance in the two regions in terms of sex ratio, age structure (n = 182 adults and yearlings), and reproductive success (n = 98 females) on the basis of 8 years of data (1995-1999, 2002-2004). Region strongly influenced females' breeding success, which, unsuccessful hens included, was consistently higher in the central region (n = 77 females) of the island than in the south (n = 21 females, P = 0.01). The central region also had a much higher proportion of successful hens (87%) than did the south (55%, P < 0.001). In light of our findings, we suggest that population performance is influenced by a combination of fine-scale habitat features and coarse-scale habitat configuration. ?? The Cooper Ornithological Society 2011.

Condor↗

Nest-site fidelity and dispersal of Gyrfalcons estimated by noninvasive genetic sampling

We used feathers from adult Gyrfalcons (Falco rusticolus) molted in breeding territories and blood samples from nestlings to document nest-site fidelity and dispersal of breeding adults and juveniles at three areas 100- 350 km apart in Yukon Delta National Wildlife Refuge, Alaska, 2003-2007. We used genotypes from seven polymorphic microsatellite loci that provided a mean probability of identity of 0.91 ??10 -5. Breeding Gyrfalcons were highly faithful to study area and territory; we documented no dispersals of breeding birds among study areas and only one dispersal between territories. But their fidelity to nest sites was low; 22% of birds returned to the same nest site the following year. Distance among alternate nests within a territory averaged 750 m and was similar for both sexes. Mean tenure in a territory was 2.8 years, similar for both sexes, and distributed bimodally with peaks at 1 and 4 years. Mean annual turnover rate at the Ingakslugwat Hills (Volcanoes) study area was 20%. We detected three young that established breeding territories at distances ranging from 0 to 254 km from their natal territory, representing 2.5% apparent recruitment. Gyrfalcons in the Askinuk Mountains study area were slightly but statistically significantly differentiated genetically from those in the Volcanoes and Kilbuck Mountain study areas. These data are the first published on the nest-site fidelity, breeding dispersal, and natal dispersal of the Gyrfalcon in North America and demonstrate the utility of noninvasive genetic sampling to greatly improve our understanding of avian dispersal and its underlying mechanisms. ?? The Cooper Ornithological Society 2011.

Conference Paper↗

Nest predation and circulating corticosterone levels within and among species

Variation in the risk of predation to offspring can influence the expression of reproductive strategies both within and among species. Appropriate expression of reproductive strategies in environments that differ in predation risk can have clear advantages for fitness. Although adult-predation risk appears to influence glucocorticosteroid levels, leading to changes in behavioral and life-history strategies, the influence of offspring-predation risk on adult glucocorticosteroid levels remains unclear. We compared total baseline corticosterone concentrations in Gray-headed Juncos (Junco hyemalis dorsalis) nesting on plots with and without experimentally reduced risk of nest predation. Despite differences in risk between treatments, we failed to find differences in total baseline corticosterone concentrations. When we examined corticosterone concentrations across a suite of sympatric species, however, higher risk of nest predation correlated with higher total baseline corticosterone levels. As found previously, total baseline corticosterone was negatively correlated with body condition and positively correlated with date of sampling. However, we also found that corticosterone levels increased seasonally, independent of stage of breeding. Nest predation can alter the expression of birds' reproductive strategies, but our findings suggest that total baseline corticosterone is not the physiological mechanism regulating these responses.

The Condor↗

A review of the population estimation approach of the North American landbird conservation plan

As part of their development of a continental plan for monitoring landbirds (Rich et al. 2004), Partners in Flight (PIF) applied a new method to make preliminary estimates of population size for all 448 species of landbirds present in the continental United States and Canada (Table 1). Estimation of the global population size of North American landbirds was intended to (1) identify the degree of vulnerability of each species, (2) provide estimates of the current population size for each species, and (3) provide a starting point for estimating population sizes in states, provinces, territories, and Bird Conservation Regions (Rich et al. 2004). A method proposed by Rosenberg and Blancher (2005) was used to derive population estimates from available survey data. To enhance the credibility of these estimates, PIF organized a review of the methodology used to estimate North American landbird population sizes. A planning committee selected members from the ornithological and biometrical communities (hereafter “the panel”), with the aim of selecting individuals from academia, state natural-resource agencies, and the U.S. and Canadian federal governments, including the Canadian Wildlife Service, the U.S. Geological Survey, and the U.S. Department of Agriculture Forest Service. The panel addressed three questions: (1) Were the methods of population estimation proposed by PIF reasonable? (2) What actions could be taken to improve the data or analyses on which the PIF population estimates were based? and (3) How should the PIF population estimates be interpreted?

The Auk↗

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.

The Auk↗

Population structure of Pacific Common Eiders breeding in Alaska

We used satellite telemetry to study the migration routes and wintering areas of two allopatric breeding populations of Pacific Common Eiders (Somateria mollissima v-nigrum) in Alaska: the Yukon-Kuskokwim Delta, and the western Beaufort Sea coast. Only 6% (2 of 36) of females wintered within the wintering area of the other breeding population. Both breeding populations wintered in the closest available ice-free habitat, perhaps to minimize migratory distance. Two Beaufort Sea females wintered in areas used by Yukon-Kuskokwim Delta females, implying potential gene flow among breeding areas. Yet, we conclude that these two populations are largely geographically isolated throughout the annual cycle and the environmental factors influencing survival and reproduction likely differ between these groups of birds. Thus, regardless of the potential gene flow among breeding populations, we suggest that birds from these two breeding areas should be managed as separate populations. ?? The Cooper Ornithological Society 2002.

Condor↗