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Migratory species can experience limiting factors at different locations and during different periods of their annual cycle. In migratory birds, these factors may even occur in different hemispheres. Therefore, identifying the distribution of populations throughout their annual cycle (i.e., migratory connectivity) can reveal the complex ecological and evolutionary relationships that link species and ecosystems across the globe and illuminate where and how limiting factors influence population trends. A growing body of literature continues to identify species that exhibit weak connectivity wherein individuals from distinct breeding areas co-occur during the nonbreeding period. A detailed account of a broadly distributed species exhibiting strong migratory connectivity in which nonbreeding isolation of populations is associated with differential population trends remains undescribed. Here, we present a range-wide assessment of the nonbreeding distribution and migratory connectivity of two broadly dispersed Nearctic-Neotropical migratory songbirds. We used geolocators to track the movements of 70 Vermivora warblers from sites spanning their breeding distribution in eastern North America and identified links between breeding populations and nonbreeding areas. Unlike blue-winged warblers (Vermivora cyanoptera), breeding populations of golden-winged warblers (Vermivora chrysoptera) exhibited strong migratory connectivity, which was associated with historical trends in breeding populations: stable for populations that winter in Central America and declining for those that winter in northern South America.

We compared the efficacy of standard nest-searching methods with finding nests via radio-tagged birds to assess how search technique influenced our determination of nest-site characteristics and nest success for Golden-winged Warblers (Vermivora chrysoptera). We also evaluated the cost-effectiveness of using radio-tagged birds to find nests. Using standard nest-searching techniques for 3 populations, we found 111 nests in locations with habitat characteristics similar to those described in previous studies: edges between forest and relatively open areas of early successional vegetation or shrubby wetlands, with 43% within 5 m of forest edge. The 83 nests found using telemetry were about half as likely (23%) to be within 5 m of forest edge. We spent little time searching >25 m into forest because published reports state that Golden-winged Warblers do not nest there. However, 14 nests found using telemetry (18%) were >25 m into forest. We modeled nest success using nest-searching method, nest age, and distance to forest edge as explanatory variables. Nest-searching method explained nest success better than nest age alone; we estimated that nests found using telemetry were 10% more likely to fledge young than nests found using standard nest-searching methods. Although radio-telemetry was more expensive than standard nest searching, the cost-effectiveness of both methods differed depending on searcher experience, amount of equipment owned, and bird population density. Our results demonstrate that telemetry can be an effective method for reducing bias in Golden-winged Warbler nest samples, can be cost competitive with standard nest-searching methods in some situations, and is likely to be a useful approach for finding nests of other forest-nesting songbirds.

Radio-transmitters and light-level geolocators are currently small enough for use on songbirds weighing <15 g. Various methods are used to attach these markers to larger songbirds, but with small birds it becomes especially important to minimize marker mass and bird handling time. Here, we offer modifications to harness materials and marker preparation for transmitters and geolocators, and we describe deployment methods that can be safely completed in 20–60 s per bird. We describe a 0.5-mm elastic sewing thread harness for radio-transmitters that allows nestlings, fledglings, and adults to be marked with the same harness size and reliably falls off to avoid poststudy effects. We also describe a 0.5-mm jewelry cord harness for geolocators that provides a firm fit for >1 yr. Neither harness type requires plastic or metal tubes, rings, or other attachment fixtures on the marker, nor do they require crimping beads, epoxy, scissors, or tying knots while handling birds. Both harnesses add 0.03 g to the mass of markers for small wood-warblers (Parulidae). This minimal additional mass is offset by trimming transmitter antennas or geolocator connection nodes, resulting in no net mass gain for transmitters and 0.02 g added for geolocators compared with conventional harness methods that add >0.40 g. We and others have used this transmitter attachment method with several small songbird species, with no effects on adult and fledgling behavior and survival. We have used this geolocator attachment method on 9-g wood-warblers with no effects on return rates, return dates, territory fidelity, and body mass. We hope that these improvements to the design and deployment of the leg-loop harness method will enable the safe and successful use of these markers, and eventually GPS and other tags, on similarly small songbirds.

ContextUnderstanding the factors limiting populations of animals is critical for effective conservation. Determining which factors limit populations of migratory species can be especially challenging because of their reliance on multiple, often geographically distant regions during their annual cycles.ObjectivesWe investigated whether distribution-wide variation in recent breeding population trends was more strongly associated with exposure to risk factors experienced during migration (i.e., natural and anthropogenic threats often associated with increased mortality or carry-over effects) or factors associated with breeding and nonbreeding areas in golden-winged warblers (Vermivora chrysoptera) and blue-winged warblers (V. cyanoptera), two Nearctic-Neotropical migrants experiencing regionally variable population trends.MethodsWe used geolocator data from 85 Vermivora warblers (n = 90 geolocator tracks) tracked from North American breeding locations and Central American nonbreeding locations from 2013 to 2017 to determine variation in space use among populations. We assessed whether differences in space use among populations of Vermivora warblers during migration were associated with exposure to migration risk-factors and whether increased relative exposure to migration risk factors was associated with population declines at regional and subregional scales.ResultsRegional and subregional populations of Vermivora warblers exhibited variation in space use and exposure to anthropogenic and natural risk-factors. However, we found no evidence that recent variation in population trends of Vermivora warblers was associated with risk-factors experienced by different populations during migration. Instead, factors associated with land cover-types in breeding and nonbreeding areas were more strongly associated with recent population trends.ConclusionsUnderstanding how populations of migratory birds are affected by factors experienced during migration is critical for their conservation. We did not find evidence that variation in exposure to migration risk-factors is associated with recent regional or subregional variation in Vermivora warbler population trends. Consequently, our results suggest that efforts to reverse ongoing population declines of Vermivora warblers may be more effective if directed toward conservation actions targeting limiting factors within the breeding and nonbreeding periods versus those directed at conditions encountered during migration. We caution that geographic variation in projected land-use change may differentially affect areas used by different populations of Vermivora warblers during migration, posing a potential threat to these species in the future.

Golden-winged Warblers (Vermivora chrysoptera) are Nearctic–Neotropical migrants experiencing varied regional population trends not fully explained by breeding-grounds factors such as nest success. A lack of detailed information on the nonbreeding distributions, migration routes, or timing of migration among populations hampers our ability to identify population processes outside the breeding period. We used geolocators to track annual movements of 21 Golden-winged Warblers from 3 North American breeding locations experiencing varying population trends to investigate the potential for nonbreeding site factors to influence breeding populations. We used the template-fit method to estimate locations of individual warblers throughout the year. Geolocator-marked warblers exhibited significant isolation among populations during migration and the nonbreeding period. During the nonbreeding period, Golden-winged Warblers from Minnesota, USA (n = 12) occurred in Central America from southern Mexico to central Nicaragua; warblers from Tennessee, USA (n = 7) occurred along the border of northern Colombia and Venezuela; and warblers from Pennsylvania, USA (n = 2) occurred in north-central Venezuela. Warblers travelled at slower rates over more days in fall migration than spring migration. Fall migration routes at the Gulf of Mexico were population-specific, whereas spring routes were more varied and overlapped among populations. Golden-winged Warblers from Pennsylvania migrated 4,000 and 5,000 km yr−1 farther than Tennessee and Minnesota warblers, respectively, and spent almost twice as long migrating in the fall compared to Minnesota warblers. Our results reveal nearly complete temporal and geographic isolation among 3 populations of Golden-winged Warblers throughout the annual cycle, resulting in opportunities for population- and site-specific factors to differentially influence populations outside the breeding period. Our findings highlight the need for monitoring multiple populations of migratory species to understand and better inform conservation strategies.

The use of light-level geolocators is increasingly common for connecting breeding and nonbreeding sites and identifying migration routes in birds. Until recently, the mass and size of geolocators precluded their use on songbird species weighing <12 g. Reducing the mass of geolocators, such as by shortening or eliminating the light stalk, may make their deployment on small birds feasible, but may also inhibit their ability to receive light reliably, because small geolocators can be shaded by feathers. Here we report geolocator effects on migratory ecology of Golden-winged Warblers (Vermivora chrysoptera) in Minnesota and Tennessee. We also evaluated whether stalk length influenced precision of location data for birds on the breeding grounds. At 8–10 g, Golden-winged Warblers are the smallest birds to be outfitted with geolocators to date. We found no differences in return rates, inter-annual territory fidelity, or body mass between geolocator-marked individuals and a control group of color-banded individuals. We observed no difference in return rates or variation in estimated breeding locations between birds marked with stalked geolocators and those with stalkless geolocators. Our results suggest that some small songbirds can be safely marked with geolocators. Light stalks appear to be unnecessary for Golden-winged Warblers; the added mass and drag of stalks can probably be eliminated on other small songbirds.

Relative nestling condition, typically measured as nestling mass or as an index including nestling mass, is commonly purported to correlate with fledgling songbird survival. However, most studies directly investigating fledgling survival have found no such relationship. We weighed feces and stomach contents of nestling golden-winged warblers (Vermivora chrysoptera) to investigate the potential contribution of variation in digestive contents to differences in nestling mass. We estimated that the mass of a seventh-day (near fledging) nestling golden-winged warbler varies by 0.65 g (approx. 9% of mean nestling mass) depending on the contents of the nestling's digestive system at the time of weighing, and that digestive contents are dissimilar among nestlings at any moment the brood is removed from the nest for weighing. Our conservative estimate of within-individual variation in digestive contents equals 72% and 24% of the mean within-brood and population-wide range in nestling mass, respectively. Based on our results, a substantive but typically unknown amount of the variation in body mass among nestlings is confounded by differences in digestive contents. We conclude that short-term variation in digestive contents likely precludes the use of body mass, and therefore any mass-dependent index, as a measure of relative nestling condition or as a predictor of survival in golden-winged warblers and likely in many other songbirds of similar size.

Investigating the potential effects of handling and marking techniques on study animals is important for correct interpretation of research results and to effect progress in data-collection methods. Few investigators have compared the reproductive output of radio-tagged and non-radio-tagged songbirds, and no one to date has examined the possible effect of radio-tagging adult songbirds on the survival of their fledglings. In 2011 and 2012, we compared several parameters of reproductive output of two groups of female Golden-winged Warblers (Vermivora chrysoptera) breeding in Minnesota, including 45 females with radio-transmitters and 73 females we did not capture, handle, or mark. We found no difference between groups in clutch sizes, hatching success, brood sizes, length of incubation and nestling stages, fledging success, number of fledglings, or survival of fledglings to independence. Thus, radio-tags had no measurable impact on the productivity of female Golden-winged Warblers. Our results build upon previous studies where investigators have reported no effects of radio-tagging on the breeding parameters of songbirds by also demonstrating no effect of radio-tagging through the post-fledging period and, therefore, the entire breeding season. Investigar los efectos potenciales de las técnicas de manipulación y marcaje en estudios de animales es importante para interpretar correctamente los resultados de las investigaciones y para llevar a cabol los avances en los métodos de colecta de datos. Pocos investigadores han comparado el rendimiento reproductivo de las aves paserinas con o sin radios trasmisores, y nadie hasta la fecha ha examinado el posible efecto en la supervivencia de los juveniles de aves marcadas con radios trasmisores. En el 2011 y 2012 comparamos varios parámetros reproductivos en dos grupos de hembras de Vermivora chrysoptera reproduciéndose en Minnesota, los cuales incluían 45 hembras con radio trasmisores y 72 hembras que no capturamos, manipulamos o marcamos. No encontramos diferencias entre los grupos en el tamaño de la nidada, éxito de eclosión, numero de polluelos, duración del periodo de incubación o polluelos, éxito de salida de los polluelos del nido o supervivencia de los juveniles hasta su independencia. En consecuencia, radio trasmisores no tienen un impacto apreciable en la productividad de hembras de V. chrysoptera. Nuestros resultados aportan ha estudios anteriores en donde investigadores no han encontrados efecto de los radio trasmisores sobre parámetros reproductivos de aves paserinas y también demuestra que no hay un efecto de los radio trasmisores sobre la supervivencia de los juveniles a lo largo del periodo después del abandono del nido, y por ende durante toda la temporada reproductiva.

Golden-winged Warblers (Vermivora chrysoptera) are Nearctic-Neotropical migrants experiencing varied regional population trends not fully explained by breeding-grounds factors such as nest success. A lack of detailed information on the nonbreeding distributions, migration routes, or timing of migration among populations hampers our ability to identify population processes outside the breeding period. We used geolocators to track annual movements of 21 Golden-winged Warblers from 3 North American breeding locations experiencing varying population trends to investigate the potential for nonbreeding site factors to influence breeding populations. We used the template-fit method to estimate locations of individual warblers throughout the year. Geolocator-marked warblers exhibited significant isolation among populations during migration and the nonbreeding period. During the nonbreeding period, Golden-winged Warblers from Minnesota, USA (n = 12) occurred in Central America from southern Mexico to central Nicaragua; warblers from Tennessee, USA (n = 7) occurred along the border of northern Colombia and Venezuela; and warblers from Pennsylvania, USA (n = 2) occurred in north-central Venezuela. Warblers travelled at slower rates over more days in fall migration than spring migration. Fall migration routes at the Gulf of Mexico were population-specific, whereas spring routes were more varied and overlapped among populations. Golden-winged Warblers from Pennsylvania migrated 4,000 and 5,000 km yr^sup -1^ farther than Tennessee and Minnesota warblers, respectively, and spent almost twice as long migrating in the fall compared to Minnesota warblers. Our results reveal nearly complete temporal and geographic isolation among 3 populations of Golden-winged Warblers throughout the annual cycle, resulting in opportunities for population- and site-specific factors to differentially influence populations outside the breeding period. Our findings highlight the need for monitoring multiple populations of migratory species to understand and better inform conservation strategies.