A neonicotinoid insecticide reduces fueling and delays migration in songbirds.

Neonicotinoids are neurotoxic insecticides widely used as seed treatments, but little is known of their effects on migrating birds that forage in agricultural areas. We tracked the migratory movements of imidacloprid-exposed songbirds at a landscape scale using a combination of experimental dosing and automated radio telemetry. Ingestion of field-realistic quantities of imidacloprid (1.2 or 3.9 milligrams per kilogram body mass) by white-crowned sparrows (Zonotrichia leucophrys) during migratory stopover caused a rapid reduction in food consumption, mass, and fat and significantly affected their probability of departure. Birds in the high-dose treatment stayed a median of 3.5 days longer at the site of capture after exposure as compared with controls, likely to regain fuel stores or recover from intoxication. Migration delays can carry over to affect survival and reproduction; thus, these results confirm a link between sublethal pesticide exposure and adverse outcomes for migratory bird populations.

Imidacloprid and chlorpyrifos insecticides impair migratory ability in a seed-eating songbird.

Birds that travel long distances between their wintering and breeding grounds may be particularly susceptible to neurotoxic insecticides, but the influence of insecticides on migration ability is poorly understood. Following acute exposure to two widely used agricultural insecticides, imidacloprid (neonicotinoid) and chlorpyrifos (organophosphate), we compared effects on body mass, migratory activity and orientation in a seed-eating bird, the white-crowned sparrow (Zonotrichia leucophrys). During spring migration, sparrows were captured, held and dosed by gavage daily for 3 days with either the vehicle control, low (10% LD50) or high (25% LD50) doses of imidacloprid or chlorpyrifos and tested in migratory orientation trials pre-exposure, post-exposure and during recovery. Control birds maintained body mass and a seasonally appropriate northward orientation throughout the experiment. Imidacloprid dosed birds exhibited significant declines in fat stores and body mass (mean loss: -17% low, -25% high dose) and failed to orient correctly. Chlorpyrifos had no overt effects on mass but significantly impaired orientation. These results suggest that wild songbirds consuming the equivalent of just four imidacloprid-treated canola seeds or eight chlorpyrifos granules per day over 3 days could suffer impaired condition, migration delays and improper migratory direction, which could lead to increased risk of mortality or lost breeding opportunity.

Ecological Causes and Consequences of Intratropical Migration in Temperate-Breeding Migratory Birds.

New discoveries from direct tracking of temperate-breeding passerines show that intratropical migration (ITM) occurs in a growing number of species, which has important implications for understanding their evolution of migration, population dynamics, and conservation needs. Our large sample size ([Formula: see text]) for purple martins (Progne subis subis) tracked with geolocators to winter sites in Brazil, combined with geolocator deployments at breeding colonies across North America, allowed us to test hypotheses for ITM, something which has not yet been possible to do for other species. ITM in purple martins was not obligate; only 44% of individuals exhibited ITM, and movements were not coordinated in time or space. We found no evidence to support the resource hypothesis; rainfall and temperature experienced by individual birds during their last 2 weeks at their first roost site were similar to conditions at their second roost site after ITM. Birds generally migrated away from the heavily forested northwestern Amazon to less forested regions to the south and east. ITM in this aerial insectivore appears to support the competition-avoidance hypothesis and may be triggered by increasing local density in the core wintering region. Full life cycle models and migratory networks will need to incorporate ITM to properly address seasonal carryover effects and identify which wintering regions are most important for conservation.

Effects of breeding versus winter habitat loss and fragmentation on the population dynamics of a migratory songbird.

Many migratory species are in decline and understanding these declines is challenging because individuals occupy widely divergent and geographically distant habitats during a single year and therefore populations across the range are interconnected in complex ways. Network modeling has been used to show, theoretically, that shifts in migratory connectivity patterns can occur in response to habitat or climate changes and that habitat loss in one region can affect sub-populations in regions that are not directly connected. Here, we use a network model, parameterized by integrating long-term monitoring data with direct tracking of -100 individuals, to explain population trends in the rapidly declining Wood Thrush (Hylocichla mustelina) and to predict future trends. Our model suggests that species-level declines in Wood Thrush are driven primarily by tropical deforestation in Central America but that protection of breeding habitat in some regions is necessary to prevent shifts in migratory connectivity and to sustain populations in all breeding regions. The model illustrates how shifts in migratory connectivity may lead to unexpected population declines in key regions. We highlight current knowledge gaps that make modeling full life-cycle population demographics in migratory species challenging but also demonstrate that modeling can inform conservation while these gaps are being filled.

Carry-Over Effects of Nonbreeding Habitat on Start-to-Finish Spring Migration Performance of a Songbird.

For migratory animals, conditions during the nonbreeding period may carry-over to influence spring migration performance. Animals in low-quality habitats are predicted to be in poorer condition, show later migration timing, and travel at slower speeds. This can result in subsequent negative effects on fitness. We tested the hypothesis that nonbreeding season body condition and habitat quality carry-over to affect spring migration performance of a long-distance migratory songbird, the Wood Thrush (Hylocichla mustelina). We tracked individual birds between multiple breeding sites in North America and nonbreeding sites in Central America. First, we compared body condition of nonbreeding birds migrating to the same general region of the breeding range with spring migration performance (timing, speed, and duration) obtained from light-level geolocators. Second, we assessed the Normalized Difference Vegetation Index (NDVI) as a proxy for nonbreeding habitat quality, and predicted that birds from wetter habitat or in wetter years (higher NDVI) would show improved migration performance relative to birds from drier sites. We found no evidence of individual-level carry-over effects of nonbreeding season body condition on spring migration performance. Lower NDVI of nonbreeding habitat resulted in delayed spring migration departure, but this effect disappeared by arrival at breeding sites. Birds occupying drier nonbreeding sites migrated faster and for fewer days, compensating for their relatively late departure. We also documented a broader pattern in NDVI and migration timing and distance, in that birds that occupied the wettest areas in the southern part of the nonbreeding range departed significantly later and migrated farther. Our results suggest that individual carry-over effects of nonbreeding habitat quality may be compensated for by a faster and shorter migration strategy. At a broad scale, consistently later spring timing and longer migration distances were associated with the wettest areas (the highest quality habitats) of the Wood Thrush non-breeding range. This supports the theory that high-quality habitats offset the costs of farther migration, resulting in a leap-frog migration pattern.

Seasonal change in tropical habitat quality and body condition for a declining migratory songbird.

Many migratory songbirds spend their non-breeding season in tropical humid forests, where climate change is predicted to increase the severity and frequency of droughts and decrease rainfall. For conservation of these songbirds, it is critical to understand how resources during the non-breeding season are affected by seasonal patterns of drying, and thereby predict potential long-term effects of climate change. We studied habitat quality for a declining tropical forest-dwelling songbird, the wood thrush (Hylocichla mustelina), and tested the hypothesis that habitat moisture and arthropod abundance are drivers of body condition during the overwintering period. We examined habitat moisture, abundance of arthropods and fruit, and condition of individual birds (n = 418) in three habitat types--mature forest, mature forest with increased presence of human activity, and riparian scrub--from October to April. We found a strong pattern of habitat drying from October (wet season) to March (prior to spring migration) in all habitats, with concurrent declines in arthropod and fruit abundance. Body condition of birds also declined (estimated ~5 % decline over the wintering period), with no significant difference by habitat. Relatively poor condition (low body condition index, low fat and pectoral muscles scores) was equally apparent in all habitat types in March. Climate change is predicted to increase the severity of dry seasons in Central America, and our results suggest that this could negatively affect the condition of individual wood thrushes.

Connectivity of wood thrush breeding, wintering, and migration sites based on range-wide tracking.

Many migratory animals are experiencing rapid population declines, but migration data with the geographic scope and resolution to quantify the complex network of movements between breeding and nonbreeding regions are often lacking. Determining the most frequently used migration routes and nonbreeding regions for a species is critical for understanding population dynamics and making effective conservation decisions. We tracked the migration of individual Wood Thrushes (Hylocichla mustelina) (n = 102) from across their range with light-level geolocators and, for the first time, quantified migration routes and wintering regions for distinct breeding populations. We identified regional and species-level migratory connectivity networks for this declining songbird by combining our tracking results with range-wide breeding abundance estimates and forest cover data. More than 50% of the species occupied the eastern wintering range (Honduras to Costa Rica), a region that includes only one-third of all wintering habitat and that is undergoing intensive deforestation. We estimated that half of all Wood Thrushes in North America migrate south through Florida in fall, whereas in spring approximately 73% funnel northward through a narrow span along the central U.S. Gulf Coast (88-93°W). Identifying migratory networks is a critical step for conservation of songbirds and we demonstrated with Wood Thrushes how it can highlight conservation hotspots for regional populations and species as a whole.

Tracking from the tropics reveals behaviour of juvenile songbirds on their first spring migration.

Juvenile songbirds on spring migration travel from tropical wintering sites to temperate breeding destinations thousands of kilometres away with no prior experience to guide them. We provide a first glimpse at the migration timing, routes, and stopover behaviour of juvenile wood thrushes (Hylocichla mustelina) on their inaugural spring migration by using miniaturized archival geolocators to track them from Central America to the U.S. and Canada. We found significant differences between the timing of juvenile migration and that of more experienced adults: juveniles not only departed later from tropical wintering sites relative to adults, they also became progressively later as they moved northward. The increasing delay was driven by more frequent short stops by juveniles along their migration route, particularly in the U.S. as they got closer to breeding sites. Surprisingly, juveniles were just as likely as adults to cross the Gulf of Mexico, an open-water crossing of 800-1000 km, and migration route at the Gulf was not significantly different for juveniles relative to adults. To determine if the later departure of juveniles was related to poor body condition in winter relative to adults, we examined percent lean body mass, fat scores, and pectoral muscle scores of juvenile versus adult birds at a wintering site in Belize. We found no age-related differences in body condition. Later migration timing of juveniles relative to adults could be an adaptive strategy (as opposed to condition-dependent) to avoid the high costs of fast migration and competition for breeding territories with experienced and larger adults. We did find significant differences in wing size between adults and juveniles, which could contribute to lower flight efficiency of juveniles and thus slower overall migration speed. We provide the first step toward understanding the "black box" of juvenile songbird migration by documenting their migration timing and en route performance.

A trans-hemispheric migratory songbird does not advance spring schedules or increase migration rate in response to record-setting temperatures at breeding sites.

The decline of long distance migratory songbirds has been linked to an increasing mismatch between spring arrival date and timing of food availability caused by climate change. It is unclear to what extent individuals can adjust migration timing or en route rate in response to annual variation in temperature at breeding sites. We tracked the ca. 7300 km spring migration of 52 purple martins Progne subis from the Amazon basin to two breeding sites in eastern North America. Spring 2012 was the warmest on record in eastern North America, but contrary to predictions, this did not result in earlier departure, faster migration, or earlier arrival at breeding areas compared with earlier years. Temperatures and rainfall in the Amazon basin at the time of departure were not higher in 2012, and conditions along migration routes did not give consistent signals of a warmer spring at the breeding site. Once in North America, individuals likely had limited opportunity to speed up their migration because this final portion of the journey was already very rapid (570 km/d; 4-5 d in duration). Migration timing over the entire journey was best predicted by breeding latitude and sex and was not sensitive to ecological cues (temperature and rainfall amount) at departure from South American overwintering sites or en route, in contrast to recent studies of other songbirds. Our results provide the first direct evidence for a mismatch between higher spring temperatures at breeding sites and departure schedules of individual songbirds, and suggest phenotypic responses to short-term climatic warming may be limited for some species. Further direct-tracking data with greater geographic and temporal scope is needed to test for individual plasticity in response to temperature and rainfall along migratory routes for this, and other, species.

Continent-wide tracking to determine migratory connectivity and tropical habitat associations of a declining aerial insectivore.

North American birds that feed on flying insects are experiencing steep population declines, particularly long-distance migratory populations in the northern breeding range. We determine, for the first time, the level of migratory connectivity across the range of a songbird using direct tracking of individuals, and test whether declining northern populations have higher exposure to agricultural landscapes at their non-breeding grounds in South America. We used light-level geolocators to track purple martins, Progne subis, originating from North American breeding populations, coast-to-coast (n = 95 individuals). We show that breeding populations of the eastern subspecies, P. s. subis, that are separated by ca. 2000 km, nevertheless have almost completely overlapping non-breeding ranges in Brazil. Most (76%) P. s. subis overwintered in northern Brazil near the Amazon River, not in the agricultural landscape of southern Brazil. Individual non-breeding sites had an average of 91 per cent forest and only 4 per cent agricultural ground cover within a 50 km radius, and birds originating from declining northern breeding populations were not more exposed to agricultural landscapes than stable southern breeding populations. Our results show that differences in wintering location and habitat do not explain recent trends in breeding population declines in this species, and instead northern populations may be constrained in their ability to respond to climate change.

Repeat tracking of individual songbirds reveals consistent migration timing but flexibility in route.

Tracking repeat migratory journeys of individual animals is required to assess phenotypic plasticity of individual migration behaviour in space and time. We used light-level geolocators to track the long-distance journeys of migratory songbirds (wood thrush, Hylocichla mustelina), and, for the first time, repeat journeys of individuals. We compare between- and within-individual variation in migration to examine flexibility of timing and route in spring and autumn. Date of departure from wintering sites in Central America, along with sex and age factors, explained most of the variation (71%) in arrival date at North American breeding sites. Spring migration showed high within-individual repeatability in timing, but not in route. In particular, spring departure dates of individuals were highly repeatable, with a mean difference between years of just 3 days. Autumn migration timing and routes were not repeatable. Our results provide novel evidence of low phenotypic plasticity in timing of spring migration, which may limit the ability of individuals to adjust migration schedules in response to climate change.

Corticosterone stress response and plasma metabolite levels during breeding and molt in a free-living migratory songbird, the wood thrush (Hylocichla mustelina).

Many birds face energetic trade-offs between different life history stages, such as reproductive effort, feather molt and the non-breeding period. Little is known about how physiological measures of condition (corticosterone, plasma metabolites) in free-living birds change from nesting stages to the post-breeding molt period or whether this is influenced by prior reproductive effort. We evaluated whether corticosterone (CORT) and plasma metabolite levels vary with date, nest stage and sex in a free-living migratory songbird, the wood thrush (Hylocichla mustelina). We also tested whether (1) baseline CORT levels early in the season were predictive of subsequent reproductive success and (2) whether prior reproductive effort influenced CORT levels and blood metabolites during molt. Baseline CORT levels decreased with date during both the incubation stage and nestling stage, but did not vary significantly across stage of breeding season. Stress-induced CORT declined with date during incubation and varied significantly across breeding stage, with lower levels during feather molt. Profiles of the metabolites of ß-hydroxybutyrate, glycerol, and triglyceride did not vary significantly with date or breeding stage. Only triglycerides varied significantly with sex, with females having higher levels than males. Reproductive output was highly variable (0-10 fledglings per season) but baseline CORT levels in females during the first incubation period of the season was not related to subsequent reproductive output. Prior reproductive effort, measured as the cumulative number of young hatched during the breeding season, was positively related to stress-induced CORT during molt. High reproductive effort in wood thrush appears to have physiological carry-over effects into the molt period which could potentially affect rate of molt and preparation for fall migration.

Effects of post-breeding moult and energetic condition on timing of songbird migration into the tropics.

Each autumn billions of songbirds migrate between the temperate zone and tropics, but little is known about how events on the breeding grounds affect migration to the tropics. Here, we use light level geolocators to track the autumn migration of wood thrushes Hylocichla mustelina and test for the first time if late moult and poor physiological condition prior to migration delays arrival on the winter territory. Late nesting thrushes postponed feather moult, and birds with less advanced moult in August were significantly farther north on 10 October while en route to the tropics. Individuals in relatively poor energetic condition in August (high ß-Hydroxybutyrate, low triglyceride, narrow feather growth bars) passed into the tropics significantly later in October. However, late moult and poor pre-migratory condition did not result in late arrival on the winter territory because stopover duration was highly variable late in migration. Although carry-over effects from the winter territory to spring migration may be strong in migratory songbirds, our study suggests that high reproductive effort late in the season does not impose time constraints that delay winter territory acquisition.

Tracking long-distance songbird migration by using geolocators.

We mapped migration routes of migratory songbirds to the Neotropics by using light-level geolocators mounted on breeding purple martins (Progne subis) and wood thrushes (Hylocichla mustelina). Wood thrushes from the same breeding population occupied winter territories within a narrow east-west band in Central America, suggesting high connectivity of breeding and wintering populations. Pace of spring migration was rapid (233 to 577 kilometers/day) except for one individual (159 kilometers/day) who took an overland route instead of crossing the Gulf of Mexico. Identifying songbird wintering areas and migration routes is critical for predicting demographic consequences of habitat loss and climate change in tropical regions.