Timing of events on the breeding grounds for five species of sympatric warblers

ABSTRACT On the breeding grounds, migratory birds have limited time to breed and molt before autumn migration. However, few studies of long‐distance migrants have examined the phenology of these events to determine what life‐history trade‐offs might result if these activities overlap. From 2000 to 2007, I used banding data to determine the timing of migration, breeding, and primary molt for Yellow Warblers (Dendroica petechia), Yellow‐rumped Warblers (D. coronata coronata), American Redstarts (Setophaga ruticilla), Ovenbirds (Seiurus aurocapilla), and Canada Warblers (Wilsonia canadensis) at a study site in Alberta, Canada. Hatching date did not differ among species (P= 0.63), with means ranging from 27 June to 3 July. All species began primary molt between 12 July and 18 July, near the expected fledging date of offspring, and therefore all species exhibited overlap between postfledging parental care and molt. The duration of primary molt ranged from 28 d for Canada Warblers to 69 d for Yellow‐rumped Warblers. Yellow Warblers, Yellow‐rumped Warblers, and American Redstarts began autumn migration having completed about 50% of their primary molt. However, Ovenbirds departed when 21% of molt was complete, and Canada Warblers departed 2 d after completing molt. For all five species of warblers, molt did not overlap with nest‐bound breeding activities. However, molt did overlap with both postfledging care and migration. This suggests that initiating migration as soon as possible is important, possibly because earlier arrival on the wintering grounds may improve access to high quality winter habitat. Overall, warblers may maximize individual fitness by combining life‐history events that result in overlapping portions of the breeding cycle, molt, and migration.     

Foraging behavior of migrant warblers in mixed‐species flocks in Venezuelan shade coffee: interspecific differences,tree species selection,and effects of drought

Shade coffee has been identified as an important habitat for Nearctic‐Neotropical migrants during the non‐breeding season, including species of conservation concern such as Cerulean Warblers (Setophaga cerulea). To better understand habitat features important for migrants in shade coffee, we studied the foraging behavior of migrants in mixed‐species flocks at six shade‐coffee farms in the Cordillera de Merida, Venezuela, in 2008–2009 and the El Niño drought year of 2009–2010. We examined interspecific differences in foraging behavior and tree species selection of three foliage‐gleaning migrants, Blackburnian (Setophaga fusca), Cerulean, and Tennessee (Oreothlypis peregrina) warblers, and aerial‐foraging American Redstarts (Setophaga ruticilla). For morphologically similar Blackburnian and Cerulean warblers, we also examined factors influencing foraging rates (attack and movement rates), capture of large prey, and maneuver/substrate type. We found that aerial‐foraging American Redstarts foraged lower, used more aerial maneuvers, showed no tree species selection, and were less likely to forage in flocks than foliage‐gleaners. Although foraging rates were similar for Blackburnian and Cerulean warblers, the three foliage‐gleaners differed in foraging height and use of maneuvers. Cerulean Warblers foraged lower than the other two species, whereas Blackburnian Warblers used the greatest proportion of woody gleans. All three foliage‐gleaners selected Inga spp. (a commonly planted shade tree in shade‐coffee farms) for foraging, and Blackburnian and Cerulean warblers captured a greater proportion of large prey in Inga spp. than in other tree species. During the drought year, Blackburnian and Cerulean warblers captured half as many large prey and used a greater proportion of woody‐gleans. We found that interactions among behavioral, floristic, and environmental drivers influenced the foraging behavior of migrants wintering in shade coffee. Our results support those of previous studies suggesting that migrants partition resources behaviorally during the non‐breeding season, that foliage‐gleaners may benefit from the presence of shade trees, especially Inga spp., in agroforestry systems, and that drought may influence the foraging behavior of foliage‐gleaning migrants, presumably due to reduced prey availability.     

Dietary opportunism,resource partitioning,and consumption of coffee berry borers by five species of migratory wood warblers (Parulidae) wintering in Jamaican shade coffee plantations

Diets reflect important ecological interactions, but are challenging to quantify for foliage‐gleaning birds. We used regurgitated stomach samples from five primarily insectivorous species of long‐distance migrant warblers (Parulidae) wintering in two moderate‐elevation shade coffee farms in Jamaica to assess both foraging opportunism and prey resource partitioning. Our results, based primarily on 6120 prey items in 80 stomach samples collected during a one‐week period in March 2000, confirm opportunism. The diets of all five warblers, including American Redstarts (Setophaga ruticilla), Black‐and‐White Warblers (Mniotilta varia), Black‐throated Blue Warblers (S. caerulescens), Northern Parulas (S. americana), and Prairie Warblers (S. discolor), overlapped strongly based on consumption of the same prey types, even many of the same prey species (4 of 10 interspecific overlaps >0.9, range = 0.74–0.97). Moreover, all five species fed on similarly small, often patchily distributed prey, including coffee berry borers (Hypothenemus hampei; Coleoptera, Curculionidae). Nonetheless, permutational multivariate analysis of variance also revealed that the diets of these species differed significantly, primarily with respect to prey mobility (winged vs. sessile); American Redstarts fed on the most mobile prey, and Northern Parulas on the least mobile prey and a relatively restricted set of prey taxa compared to the other four species of warblers. Overall, our results suggest both dietary opportunism consistent with a migratory life‐history, and interspecific resource partitioning consistent with differences in morphology and foraging behavior during a food‐limited season. Having provided evidence of the three necessary conditions, namely intraspecific competition, resource limitation, and interspecific overlap in resource use, the results of our study, in combination with those of other studies, also provide evidence of interspecific competition among wintering migrant insectivores. We thus argue that diffuse interspecific exploitative food competition may be more important than previously recognized.     

Estimating mass change of migrant songbirds during stopover: comparison of three different methods

ABSTRACT Stopover‐site quality has often been assessed using changes in the body mass of migrants estimated from individuals recaptured on subsequent days or using regression methods. We compared estimates of mass change using these two techniques to estimates of mass change determined from birds recaptured on the same day. Using spring and fall banding data collected on Appledore Island, Maine, from 1990–2007, we examined body mass changes of the five most common species. Over this period, 18,954 individuals of these five species were captured and banded, with 11.6% of birds recaptured at least 1 d after initial capture and 3.1% recaptured on the same day. Using both regression and same‐day recapture methods, all five species had positive hourly mass gains during fall migration; results were mixed for the subsequent‐day analysis method. Trends were less consistent during spring migration. Using all three methods of estimating mass change, Red‐eyed Vireos (Vireo olivaceus) lost mass, American Redstarts (Setophaga ruticilla) and Northern Waterthrushes (Parkesia noveboracensis) gained mass, and results for Yellow‐bellied Flycatchers (Empidonax flaviventris), and Black‐and‐white Warblers (Mniotilta varia) varied with method. We found similar trends in mass change using the same‐day recapture and regression methods. However, we found lower mean mass gain for most species using the same‐day recapture method, suggesting that there may be a short‐term capture and handling effect. Our results provide additional support for the use of regression models to compare changes in mass of migrating songbirds at stopover sites.     

Cerulean Warblers in the Ozark region: habitat selection,breeding biology,survival, and space use

Cerulean Warblers (Setophaga cerulea) are a species with declining populations that exhibit regional variation in habitat selection and demographic rates. The Ozark region of the south‐central United States likely provides important habitat for Cerulean Warblers, but little is known about their breeding biology in that region. We studied Cerulean Warblers in riparian forests of the Ozarks of Arkansas from 2018 to 2020. We assessed multi‐scale habitat selection for vegetative and topographic features, documented their breeding biology, estimated within‐season and annual apparent survival, and estimated territory sizes. We found that Cerulean Warblers selected riparian habitat characterized by large‐diameter trees across all spatial scales. Contrary to the results of previous studies, males appeared to avoid white oaks (Quercus spp., Section Quercus) at the territory scale, but this avoidance may reflect an underlying preference for riparian habitat. Our logistic‐exposure estimate of nest survival (0.32; 85% confidence interval: 0.21–0.46) was similar to the median of estimates reported in previous studies. Our results indicate that maintaining riparian forests with large trees is important to provide suitable habitat for Cerulean Warblers in the Ozark region. Because of similarities in habitat selection among regions, some management practices from other populations, including retaining large trees and promoting a heterogeneous canopy structure, may be useful for managing for Cerulean Warblers in riparian areas of the Ozarks. However, selection for topography and tree species by Cerulean Warblers in our study also suggests that region‐specific management strategies will be beneficial. Finally, our demographic rate estimates for this population should prove valuable in future full‐annual‐cycle population modeling efforts.     

Influence of fine‐scale habitat requirements and riparian degradation on the distribution of the purple‐crowned fairy‐wren (Malurus coronatus coronatus) in northern Australia

Species distributions are influenced by variation in environmental conditions across many scales. Knowledge of fine‐scale habitat requirements is important for predicting species occurrence and identifying suitable habitat for target species. Here we investigate the perplexing distribution of a riparian habitat specialist, the western subspecies of the purple‐crowned fairy‐wren (Malurus coronatus coronatus), in relation to fine‐scale habitat associations and patterns of riparian degradation. Surveys of vegetation attributes, river structure and disturbance indicators that are likely to be causal determinants of the species occurrence were undertaken at 635 sites across 14 catchments. Generalized Linear Mixed Modelling demonstrated that the probability of purple‐crowned fairy‐wren occurrence increased with Pandanus aquaticus crown cover, shrub density and height of emergent trees, while riparian structure and signs of cattle were indirect predictors of occurrence. As our study area predominantly contained Pandanus type habitat, we failed to identify river grass as an important component of habitat. Predictions from a cross‐validated model of purple‐crowned fairy‐wren occurrence suggested distribution is constrained by three factors: (i) low quality of local habitat within catchments where the species occurs; (ii) broad‐scale reduction in habitat quality that has resulted in extinction of the species from parts of its range; and (iii) unmeasured variables that limit the exploitation of suitable habitat. The reliance of the species on dense shrubby understorey suggests conservation efforts should aim to maintain the complexity of understorey structure by managing fire and grazing intensity. Efforts to halt the continuing decline of riparian condition and maintain connectivity between areas of quality habitat will help to ensure persistence of riparian habitat specialists in northern Australia.     

Characteristics of the breeding habitat of Endangered Kirtland's Warblers on a Canadian Military Installation

Kirtland's Warblers (Setophaga kirtlandii) are an endangered species with specialized habitat requirements, and the only documented nesting location in Canada is an Army installation. From 2007 to 2010, I compared habitat characteristics of sites occupied and not occupied by Kirtland's Warblers at Garrison Petawawa located ～200 km northwest of Ottawa, Ontario. Sites occupied by Kirtland's Warblers (N = 11) had greater percent cover of low sweet blueberry (Vaccinium angustifolium; 31.2%), coral lichen (Cladina stellaris; 0.4%), reindeer lichen (Cladonia rangiferina; 8.3%), and wavy‐leaved moss (Dicranum polysetum; 4.3%) than unoccupied sites (N = 6). I found no difference in tree species composition between sites, with jack pine (Pinus banksiana), white pine (Pinus strobus), red pine (Pinus resinosa), poplar (Populus tremuloides), and red maple (Acer rubra) present in both used and unused sites. Forest stands occupied by Kirtland's Warblers were significantly younger (< 20 yr old) than unoccupied sites, and most occupied sites were within former fire zones and sites where jack pines had been planted. Thus, breeding habitat of Kirtland's Warblers in Canada consisted of young pine trees, with more red pine than in their breeding habitat in Michigan, and ground cover including blueberry, lichens, and mosses. These results suggest that Kirtland's Warblers may be less selective in their habitat requirements than previously thought, and should provide guidance for recovery groups and regulatory agencies in accurately delineating suitable habitat for these warblers in Canada.     

Foraging behavior of three species of songbirds during stopover in southeastern Morocco during spring migration

Investigators studying the stopover ecology of migrating birds typically use the capture–recapture method to examine important parameters such as fuel deposition rates (FDR) and stopover duration. However, such studies can be constrained by the number of recaptures. An alternative method is to calculate a regression of mass over time of day, but this method may not be reliable because patterns of mass change of individual birds through the day may not reflect that of the whole population. Given the potential constraints of these methods, using them in combination with other methods, such as behavioral observations of foraging birds, may improve our understanding of the patterns of fuelling in birds at stopover sites. We observed the foraging behavior of three songbird species, including Western Bonelli's (Phylloscopus bonelli), Subalpine (Sylvia cantillans), and Willow (Phylloscopus trochilus) warblers, from 15 March to 30 April 2011 at a small oasis at the northern border of the Sahara desert in southeast Morocco. Given the location of our study site at the northern edge of the Sahara desert, birds migrating north likely needed to replenish their energy reserves at this stage of their journey. We assessed foraging effort by determining the rate (number per unit time) at which birds pecked at substrates or made aerial forays after flying insects. Peck rates were higher for Western Bonelli's Warblers than for Subalpine and Willow warblers, suggesting either species‐specific adaptations to feeding in arid environments or differences in the motivation to feed. In addition, Western Bonelli's Warblers had FDRs that were negative or close to zero and, therefore, were apparently unable to refuel successfully (i.e., increase their fuel stores) despite greater effort, possibly indicating less efficiency in obtaining food (i.e., more unsuccessful pecks). The lower peck rates of Subalpine and Willow warblers suggest either that they were less efficient at finding prey or were simply foraging at lower rates. For all three species, peck rates were lower at higher wind speeds, suggesting that wind may alter prey availability and detectability, especially of flying insects. Interactions among species‐specific migration strategies, environmental conditions, and habitat quality ultimately define the success of migration. Our results suggest that using observational data in combination with capture data may improve our understanding of these interactions at migration stopover sites.     

Carry‐over effects of winter habitat vary with age and sex in yellow warblers Setophaga petechia

We use stable isotope data to investigate the role of winter habitat use in altering the breeding phenology of yellow warblers Setophaga petechia. We first confirm that δ¹³C and δ¹⁵N isotopic signatures vary with winter habitat use in this species. We then examine the relationship between winter habitat use, breeding phenology and productivity within four age‐sex‐classes, since life history theory would predict that carry‐over effects should vary with age and gender. The δ¹³C signatures of yellow warblers using riparian habitats over winter were more depleted than the signatures of those using agricultural or scrub habitat. Individuals on the Pacific coast of Mexico were also more δ¹⁵N enriched than those on the southern Gulf of Mexico. δ¹³C and δ¹⁵N signatures were only correlated with earlier clutch initiation and subsequent higher productivity in first‐breeding‐season females. We estimate that shifts in δ¹³C equivalent to a shift from scrub to riparian winter habitat would be associated with the production of 0.8 more fledglings by yearling females. Pre‐breeding events that influence the timing of breeding could also influence the reproductive performance of older males and females, but we found little evidence that winter habitat use influenced breeding season phenology in these birds.     

WEIGHTS OF SOME PALAEARCTIC MIGRANTS IN SOUTHERN UGANDA

Between March 1966 and May 1968 Palaearctic passerines were mist-netted in thick bush and lightly wooded savannah habitats near Kampala, on the northern shore of Lake Victoria. This paper reports weights of the seven principal species involved. Most migrants appeared to be in a lean condition during the winter months, when weights were relatively low and varied little in each species. Birds were not particularly light on arrival. In fact, autumn Garden Warblers Sylvia borin and Willow Warblers Phylloscopus trochilus were sometimes markedly heavy, and for the former species there was some evidence that the individuals concerned were passage migrants. Autumn weights of Swallows Hirundo rustica, Reed Warblers Acrocephalus scirpaceus and Yellow Wagtails Motacilla flava were similar to those recorded in winter. The mean weight of all species rose during late March or early April. Although most Garden Warblers and Willow Warblers trapped at the time of spring migration were within the normal winter weight range, many Acrocephalus warblers and the majority of Sand Martins Riparia riparia and Yellow Wagtails were rather heavy. Spring weights 40% or more above mean winter weight were not uncommon in the Sedge Warbler Acrocephalus schoenobaenus, but were recorded only occasionally in other species. Although most passerine migrants evidently left Kampala with substantial fat reserves, it was concluded that a considerable number of warblers departed at rather low weight. High spring weights were mainly confined to a period of two or three weeks in each of the warbler species. Locally wintering Acrocephalus warblers must have attained full premigratory weights within three weeks, and a number of spring retraps showed substantial gains at minimum mean rates of between 0–1 and 0–35 g per day. Most heavy Garden Warblers were probably on passage. Significant correlations between weight and wing-length were obtained for all species investigated, regressions of weight on wing-length being in the range 011-0-25 g/mm. Spring weights are briefly compared with data from Nigeria, and the northward migration of passerines from Lake Victoria is discussed.     

Detection probability of Golden-winged Warblers during point counts with and without playback recordings

ABSTRACT Use of point‐count data to estimate population sizes of North American landbirds may be challenged by limitations on detection probability of particular species, thereby requiring correction factors to ensure accurate estimates. We estimated detection probability of Golden‐winged Warblers (Vermivora chrysoptera) during 3‐min point‐count surveys conducted both with and without use of playback recordings in a mixed shrubland‐forest habitat (clearcut area) and a 60‐m wide electric transmission line right‐of‐way (ROW) in central Pennsylvania from 20 May to 17 June 2002–2003. In addition, we assessed the value of playback with respect to response rates of warblers and distance within which warblers approached the observer. Without playback, detection probability was approximately 23% in the clearcut area and 61% in the ROW. Use of playback resulted in 7% and 19% net increases in probability at the clearcut area and the ROW, respectively; proportional increase was approximately 30% for both habitats. Warblers responded to playback 68% of the time, but response rate was greater within 100 m (72%) than beyond (53%). Most responses (85%) included approach of the warbler toward the observer, and most individuals approached within 10 m. We conclude that 3‐min point counts with playback do not yield detection probabilities sufficient to estimate population size of Golden‐winged Warblers without use of correction factors. Furthermore, detection probability in mixed shrubland‐forest habitats can be much lower than in linear habitats such as utility ROWs. Efforts to estimate population size of Golden‐winged Warblers from data of the North American Breeding Bird Survey should recognize that habitat structure has much influence on detection probability as it relates to distance at which an observer can hear (or see) warblers. Accordingly, we recommend that such efforts incorporate a maximum detection distance of 100–150 m in mixed shrubland‐forest habitats.     

Habitat use and densities of co‐existing migrant Willow Warblers Phylloscopus trochilus and resident eremomelas Eremomela spp. in Zimbabwe

Capsule Migrant Willow Warblers occupy more woodland types and occur at higher densities than ecologically‐similar resident Afrotropical warblers.

Aims To compare population densities of Willow Warblers and eremomelas in adjacent acacia, mopane and miombo woodlands, and assess the abundance of potential invertebrate prey in each habitat type, in order to investigate whether Palearctic migrants use more open habitats and are more flexible in habitat use than their Afrotropical counterparts in the same feeding guild.

Methods Using distance sampling we carried out four replicated sets of point counts in acacia woodland and three sets of counts in miombo and mopane between December 1999 and February 2000. We noted the tree species in which we saw warblers foraging and took beating‐tray samples of potential arthropod prey present on tree foliage in each of the three habitats.

Results Willow Warbler density in acacia woodland increased from 1.80 ± 0.54 (se) birds/ha in early December to 7.15 ± 1.41 birds/ha in late January after influxes of later arrivals. Densities of Willow Warblers in miombo and mopane were much lower (1.14 ± 0.28 and 0.38 ± 0.23 birds/ha, respectively) and did not show significant changes. Burnt‐necked Eremomelas averaged 0.74 ± 0.34 birds/ha in acacia woodland, and in miombo densities of Green‐capped and Yellow‐bellied Eremomelas were 0.23 ± 0.17 and 0.34 ± 0.26 birds/ha, respectively. Densities in mopane were too low to estimate reliably. Willow Warblers and Green‐capped Eremomelas showed some apparent preferences in tree species used for foraging but differences in tree use were not obviously related to the abundance of arthropod taxa present as potential prey.

Conclusion Willow Warblers occupied more habitats at greater density than similar Afrotropical warblers. They appear to favour acacia, but their settlement patterns and the reasons for disparities between densities of immigrants and residents are unclear.     

Importance of Montane Riparian Forest and Influence of Wildfire on Nest-Site Selection of Ground-Nesting Birds

Abstract: We studied breeding populations of 2 coexisting ground-nesting birds, the red-faced warbler (Cardellina rubrifrons) and yellow-eyed junco (Junco phaeonotus), in high-elevation (>2,000 m) forested drainages of the Santa Catalina Mountains, Arizona, USA. From 2004 to 2005, we 1) estimated density and nesting success of breeding populations of red-faced warblers and yellow-eyed juncos, 2) identified nest-site characteristics for each species (i.e., used sites vs. random plots), 3) compared nest-site characteristics between the 2 species, and 4) examined effects of a recent (2003) wildfire on distribution of nests of both species. In addition, we estimated the areal extent of montane riparian forest (the preferred breeding habitat of both species) within high-elevation forests of the Santa Catalina Mountains. We found that red-faced warblers and yellow-eyed juncos were the 2 most common ground-nesting birds within our study area with an average density of 2.4 and 1.4 singing males/ha, respectively, along drainage bottoms. Compared to random plots, most red-faced warbler and yellow-eyed junco nests were located close (≤30 m) to drainage bottoms within a strip of montane riparian forest characterized by abundant brush, small woody debris, ferns, and forbs (both species), high number and diversity of saplings and small trees (red-faced warblers), and abundant shrubs and downed logs and less canopy cover (yellow-eyed juncos). Although both species nested in close proximity within montane riparian forest, nest-site characteristics differed between the 2 species, especially at finer spatial scales. For example, most yellow-eyed juncos nested adjacent to grass (principally Muhlenbergia spp.), whereas red-faced warblers nested adjacent to a variety of plant species, including grass, bigtooth maple (Acer grandidentatum), white fir (Abies concolor), and Douglas-fir (Pseudotsuga menziesii). Both red-faced warblers and yellow-eyed juncos avoided nesting in areas burned during a recent wildfire. In addition, nesting success was low for red-faced warblers (13%) and yellow-eyed juncos (19%) following the wildfire, suggesting an indirect negative effect of fire on breeding populations in the short-term. Montane riparian forest appears to provide important breeding habitat for red-faced warblers and yellow-eyed juncos. However, little research or conservation planning has been directed toward montane riparian forest in the region, even though this forest type is limited in its areal extent (<11% of high-elevation forest in the Santa Catalina Mountains) and increasingly threatened by disturbance. Results from our study can be used to facilitate the management and conservation of breeding populations of red-faced warblers and yellow-eyed juncos in high-elevation forests of the southwestern United States.     

Compensatory frugivory in migratory Sylvia warblers: geographical responses to season length

Seasonal shifts in the diets of Garden Warblers Sylvia borin , Blackcaps S. atricapilla and Lesser Whitethroats S. curruca at a North German site were studied from samples obtained by flushing the digestive tract of the birds. The significance of seven species of berries and pulpy fruits in the diet of these species while feeding in riparian vegetation during the premigratory and migratory seasons was assessed and compared with the pattern of frugivory shown by the same species in southern Germany. Because of the short time span between hatching and onset of migration, first-year long- and medium-distance migratory Sylvia warblers from northern Europe are forced to accelerate their juvenile development. Presumably to meet the higher nutritional demands of an accelerated development, juvenile Garden Warblers and Blackcaps in northern Germany ate relatively more fruit during premigration than Blackcaps in southern Germany. Fruit occurred in about 90% of their diet samples during both premigration and migration. However, this might not hold true for experienced adult birds. Garden Warblers accelerated the rate of fattening and reduced the time needed to prepare for migration when feeding on a mixed fruit diet. There were indications that Blackcaps directly influenced the fruit choice of Garden Warblers when fruits preferred by both species were scarce. Blackcaps and Garden Warblers showed subtle differences in their seasonal preferences for black elder Sambucus nigra and woody nightshade Solanum dulcamara .     

Carry‐over effects of winter habitat quality on en route timing and condition of a migratory passerine during spring migration

We examined how conditions prior to migration influenced migration performance of two breeding populations of black‐and‐white warblers Mniotilta varia by linking information on the migrant's winter habitat quality, measured via stable carbon isotopes, with information on their breeding destination, measured via stable hydrogen isotopes. The quality of winter habitat strongly influenced the timing of migration when we accounted for differential timing of migration between breeding populations. Among birds migrating to the same breeding destination, males and females arriving early to the stopover site originated from more mesic habitat than later arriving birds, suggesting that the benefits of occupying high‐quality mesic habitat during the winter positively influence the timing of migration. However, male warblers arriving early to the stopover site were not in better migratory condition than later arriving conspecifics that originated from poor‐quality xeric winter habitat, regardless of breeding destination. The two breeding populations stopover at the study site during different time periods, suggesting that the lower migratory condition of early birds is not a function of the time of season, but potentially a migrant's migration strategy. Strong selection pressures to arrive early on the breeding grounds to secure high‐quality breeding territories may drive males from high‐quality winter habitat to minimize time at the expense of energy. This migration strategy would result in a smaller margin of safety to buffer the effects of adverse weather or scarcity of food, increasing the risk of mortality. The migratory condition of females was the same regardless of the timing of migration or breeding destination, suggesting that females adopt a strategy that conserves energy during migration. This study fills an important gap in our understanding of the linkages between winter habitat quality and factors that influence the performance of migration, the phase of the annual cycle thought to be limiting most migratory bird populations.     

An improved survey method for monitoring population trends of Golden‐winged Warblers and other patchily distributed birds

Conventional surveys designed to monitor common and widespread species may fail to adequately track population changes of rare or patchily distributed species that are often of high conservation concern. We evaluated the performance of a new monitoring approach that employs both a spatially balanced sampling design and a targeted survey protocol designed to estimate population trends of one such patchily distributed species, the Golden‐winged Warbler (Vermivora chrysoptera), in the Appalachian Mountains Bird Conservation Region (BCR 28), USA. Our spatially balanced survey consisted of 105 sample quads (one‐quarter Delorme Atlas pages) across the current range of Golden‐winged Warblers within BCR 28, each with five sample points located in early successional habitat. From 2009 to 2013, collaborators visited each sample point once per year during the peak breeding season and conducted a 17‐min survey consisting of passive observation and playback of conspecific songs and mobbing vocalizations. We used multi‐season, single‐species occupancy models to estimate probability of quad occupancy, detection probability, and occupancy dynamics for Golden‐winged Warblers and closely related Blue‐winged Warblers (Vermivora cyanoptera). Our survey protocol resulted in high estimates of detection probability for Golden‐winged (92%) and Blue‐winged (79%) warblers, with 47% and 56% of quads estimated to be initially occupied, respectively. Derived population trend estimates (λ) indicated an average decline in population of 6% for Golden‐winged Warblers and 7% for Blue‐winged Warblers, resulting in estimated 21% and 22% declines, respectively, in quad occupancy after 5 yr. Our results demonstrate that coupling a spatially balanced survey design in appropriate habitat with a playback protocol to increase detection rates is a viable strategy for tracking populations of Golden‐winged Warblers in the Appalachian Mountains BCR. Similar survey methods should be considered for other rare, declining, or patchily distributed bird species that require targeted monitoring.     

Nesting Ecology of Boreal Forest Birds Following a Massive Outbreak of Spruce Beetles

Abstract: We studied breeding dark-eyed juncos (Junco hyemalis), yellow-rumped warblers (Dendroica coronata), and spruce-nesting birds from 1997 to 1998 among forests with different levels of spruce (Picea spp.) mortality following an outbreak of spruce beetles (Dendroctonus rufipennis) in Alaska, USA. We identified species using live and beetle-killed spruce for nest sites and monitored nests to determine how the outbreak influenced avian habitat selection and reproduction. We tested predictions that 1) nesting success of ground-nesting juncos would increase with spruce mortality due to proliferation of understory vegetation available to conceal nests from predators, 2) nesting success of canopy-nesting warblers would decrease with spruce mortality due to fewer live spruce in which to conceal nests, and 3) both species would alter nest-site selection in response to disturbance. Juncos did not benefit from changes in understory vegetation; nesting success in highly disturbed stands (46%) was comparable to that in undisturbed habitats throughout their range. In stands with low spruce mortality, nesting success of juncos was low (5%) and corresponded with high densities of red squirrels (Tamiasciurus hudsonicus). Yellow-rumped warblers nested exclusively in spruce, but success did not vary with spruce mortality. As disturbance increased, nesting warblers switched from selecting forest patches with high densities of live white spruce (Picea glauca) to patches with beetle-killed spruce. Warblers also placed nests in large-diameter live or beetle-killed spruce, depending on which was more abundant in the stand, with no differences in nesting success. Five of the 12 other species of spruce-nesting birds also used beetle-killed spruce as nest sites. Because beetle-killed spruce can remain standing for >50 years, even highly disturbed stands provide an important breeding resource for boreal forest birds. We recommend that boreal forest managers preserve uncut blocks of infested forest within managed forest landscapes and practice partial harvest of beetle-killed spruce rather than commercial clear-cutting of infested stands in order to sustain breeding bird populations until natural reforestation occurs. Because breeding densities do not always reflect fitness, assessing impacts of a massive natural disturbance should include measuring impacts of changes in vegetation on both reproductive success and predator-prey dynamics.     

Rufous‐capped Warblers Basileuterus rufifrons show seasonal,temporal and annual variation in song use

In the majority of songbird species, males have repertoires of multiple song types used for mate attraction and territory defence. The wood‐warblers (family Parulidae) are a diverse family of songbirds in which males of many migratory species use different song types or patterns of song delivery (known as ‘singing modes’) depending on context. The vocal behaviour of most tropical resident warblers remains undescribed, although these species differ ecologically and behaviourally from migratory species, and may therefore differ in their vocal behaviour. We test whether male Rufous‐capped Warblers Basileuterus rufifrons use distinct singing modes by examining song structure and context‐dependent variation in their songs. We recorded multiple song bouts from 50 male Warblers in a Costa Rican population over 3 years to describe seasonal, diel and annual variation in song structure and vocal behaviour. We found that Rufous‐capped Warbler songs are complex, with many syllable types shared both within and between males’ repertoires. Males varied their song output depending on context: they sang long songs at a high rate at dawn and during the breeding season, and shortened songs in the presence of a vocalizing female mate. Unlike many migratory species, Rufous‐capped Warblers do not appear to have different singing modes; they did not change the song variants used or the pattern of song delivery according to time of day, season or female vocal activity. Our research provides the first detailed vocal analysis of any Basileuterus warbler species, and enhances our understanding of the evolution of repertoire specialization in tropical resident songbirds.     

An ‘ecological trap’ for yellow warbler nest microhabitat selection

Contrary to assumptions of habitat selection theory, field studies frequently detect ‘ecological traps’, where animals prefer habitats conferring lower fitness than available alternatives. Evidence for traps includes cases where birds prefer breeding habitats associated with relatively high nest predation rates despite the importance of nest survival to avian fitness. Because birds select breeding habitat at multiple spatial scales, the processes underlying traps for birds are likely scale‐dependent. We studied a potential ecological trap for a population of yellow warblers Dendroica petechia while paying specific attention to spatial scale. We quantified nest microhabitat preference by comparing nest‐ versus random‐site microhabitat structure and related preferred microhabitat features with nest survival. Over a nine‐year study period and three study sites, we found a consistently negative relationship between preferred microhabitat patches and nest survival rates. Data from experimental nests described a similar relationship, corroborating the apparent positive relationship between preferred microhabitat and nest predation. As do other songbirds, yellow warblers select breeding habitat in at least two steps at two spatial scales; (1) they select territories at a coarser spatial scale and (2) nest microhabitats at a finer scale from within individual territories. By comparing nest versus random sites within territories, we showed that maladaptive nest microhabitat preferences arose during within‐territory nest site selection (step 2). Furthermore, nest predation rates varied at a fine enough scale to provide individual yellow warblers with lower‐predation alternatives to preferred microhabitats. Given these results, tradeoffs between nest survival and other fitness components are unlikely since fitness components other than nest survival are probably more relevant to territory‐scale habitat selection. Instead, exchanges of individuals among populations facing different predation regimes, the recent proliferation of the parasitic brown‐headed cowbird Molothrus ater, and/or anthropogenic changes to riparian vegetation structure are more likely explanations.