Ticks without borders: Microbial communities of immature Neotropical tick species parasitizing migratory landbirds along northern Gulf of Mexico.

The long-distance, seasonal migrations of birds make them an effective ecological bridge for the movement of ticks. The introduction of exotic tick species to new geographical regions can lead to the emergence of novel tick-borne pathogens or the re-emergence of previously eradicated ones. This study assessed the prevalence of exotic tick species parasitizing resident, short-distance, and long-distance songbirds during spring and autumn at stopover sites in the northern Gulf of Mexico using the mitochondrial 12S rDNA gene. Birds were captured for tick collection from six different sites from late August to early November in both 2018 and 2019. The highest number of ticks were collected in the 2019 season. Most ticks were collected off the Yellow-breasted Chat (Icteria virens) and Common Yellowthroat (Geothlypis trichas), and 54% of the total ticks collected were from Grand Chenier, LA. A high throughput 16S ribosomal RNA sequencing approach was followed to characterize the microbial communities and identify pathogenic microbes in all tick samples. Tick microbial communities, diversity, and community structure were determined using quantitative insight into microbial ecology (QIIME). The sparse correlations for compositional data (SparCC) approach was then used to construct microbial network maps and infer microbial correlations. A total of 421 individual ticks in the genera Amblyomma, Haemaphysalis, and Ixodes were recorded from 28 songbird species, of which Amblyomma and Amblyomma longirostre was the most abundant tick genus and species, respectively. Microbial profiles showed that Proteobacteria was the most abundant phylum. The most abundant bacteria include the pathogenic Rickettsia and endosymbiont Francisella, Candidatus Midichloria, and Spiroplasma. BLAST analysis and phylogenetic reconstruction of the Rickettsia sequences revealed the highest similarities to pathogenic spotted and non-spotted fever groups, including R. buchneri, R. conorii, R. prowazekii, R. bellii, R. australis, R. parkeri, R. monacensis, and R. monteiroi. Permutation multivariate analysis of variance revealed that the relative abundance of Francisella and Rickettsia drives microbial patterns across the tick genera. We also observed a higher percentage of positive correlations in microbe-microbe interactions among members of the microbial communities. Network analysis suggested a negative correlation between a) Francisella and Rickettsia and, b) Francisella and Cutibacterium. Lastly, mapping the distributions of bird species parasitized during spring migrations highlighted geographic hotspots where migratory songbirds could disperse ticks and their pathogens at stopover sites or upon arrival to their breeding grounds, the latter showing means dispersal distances from 421-5003 kilometers. These findings strongly highlight the potential role of migratory birds in the epidemiology of tick-borne pathogens.

Estimating apparent survival of songbirds crossing the Gulf of Mexico during autumn migration.

Many migratory bird species are declining, and the migratory period may limit populations because of the risk in traversing large geographical features during passage. Using automated radio-telemetry, we tracked 139 Swainson's thrushes (Catharus ustulatus) departing coastal Alabama, USA and crossing the Gulf of Mexico to arrive in the Yucatan Peninsula, Mexico during autumn. We estimated apparent survival and examined how extrinsic (weather variables and day of year) and intrinsic (fat load, sex and age) factors influenced survival using a mark-recapture approach. We also examined how favourability of winds for crossing the Gulf varied over the past 25 years. Fat load, day of year and wind profit were important factors in predicting which individuals survived crossing the Gulf. Survival estimates varied with wind profit and fat, but generally, fat birds departing on days with favourable wind profits had an apparent survival probability of greater than 0.90, while lean individuals with no or negative wind profits had less than 0.33. The proportion of favourable nights varied within and among years, but has increased over the last 25 years. While conservation strategies cannot improve extrinsic factors, they can provide opportunities for birds to refuel before crossing large geographical features through protecting and creating high-quality stopover sites.

Temporal migration patterns between natal locations of ruby-throated hummingbirds (Archilochus colubris) and their Gulf Coast stopover site.

BACKGROUND: Autumn latitudinal migrations generally exhibit one of two different temporal migration patterns: type 1 where southern populations migrate south before northern populations, or type 2 where northern populations overtake southern populations en route. The ruby-throated hummingbird (Archilochus colubris) is a species with an expansive breeding range, which allows opportunities to examine variation in the timing of migration. Our objective was to determine a relationship between natal origin of ruby-throated hummingbirds and arrival at a Gulf coast stopover site; and if so, what factors, such as differences in body size across the range as well as the cost of migration, might drive such a pattern. To carry out our objectives, we captured hummingbirds at a coastal stopover site during autumn migration, at which time we collected feathers from juveniles for analysis of hydrogen stable isotopes. Using the hydrogen stable isotope gradient of precipitation across North America and published hydrogen isotope values of feathers from populations of breeding ruby-throated hummingbirds, we assigned migrants to probable natal latitudes. RESULTS: Our results confirm that individuals from across the range (30-50° N) stopover along the Gulf of Mexico and there is a positive relationship between arrival day and latitude, suggesting a type 1 migration pattern. We also found no relationship between fuel load (proxy for migration cost) or fat-free body mass (proxy for body size) and natal latitude. CONCLUSIONS: Our results, coupled with previous work on the spatial migration patterns of hummingbirds, show a type 1 chain migration pattern. While the mechanisms we tested do not seem to influence the evolution of migratory patterns, other factors such as resource availability may play a prominent role in the evolution of this migration system.

Sex-specific hypothalamic-pituitary-gonadal axis sensitivity in migrating songbirds.

In seasonally migratory species, the overlap between the migratory and breeding life history stages is a balance between the physiological and behavioral requirements of each stage. Previous studies investigating the degree to which songbirds prepare for breeding during spring migration have focused on either circulating hormone levels or direct measures of gonadal recrudescence. In this study, we evaluated the phenology of breeding preparation in a long-distance migratory songbird, the Swainson's Thrush (Catharus ustulatus), by assessing hypothalamic-pituitary-gonadal (HPG) axis sensitivity with gonadotropin-releasing hormone (GnRH) bioassays throughout the migratory period. During spring migration both males and females had a significant response to GnRH injections as reflected in elevated testosterone levels. The magnitude of response to GnRH injections, Rpotential, in females stayed consistent throughout spring migration; however, Rpotential in males increased as the migratory season progressed. It is clear that at least some degree of endocrinological breeding development occurs either before or during spring migration in both sexes, however the phenology appears to be sex specific. In males this breeding development continues at a relatively steady pace throughout the migratory period while in females, relatively little endocrine breeding development occurs during migration. These sex-specific differences in the phenology of the endocrine breeding development warrant future investigations for both male and female songbirds. Moreover, research focused on how physiological breeding development is balanced with the expression of migratory traits in long-distance songbird migrants is needed.

Swainson's Thrushes do not show strong wind selectivity prior to crossing the Gulf of Mexico.

During long-distance fall migrations, nocturnally migrating Swainson's Thrushes often stop on the northern Gulf of Mexico coast before flying across the Gulf. To minimize energetic costs, trans-Gulf migrants should stop over when they encounter crosswinds or headwinds, and depart with supportive tailwinds. However, time constrained migrants should be less selective, balancing costs of headwinds with benefits of continuing their migrations. To test the hypotheses that birds select supportive winds and that selectivity is mediated by seasonal time constraints, we examined whether local winds affected Swainson's Thrushes' arrival and departure at Ft. Morgan, Alabama, USA at annual, seasonal, and nightly time scales. Additionally, migrants could benefit from forecasting future wind conditions, crossing on nights when winds are consistently supportive across the Gulf, thereby avoiding the potentially lethal consequences of depleting their energetic reserves over water. To test whether birds forecast, we developed a movement model, calculated to what extent departure winds were predictive of future Gulf winds, and tested whether birds responded to predictability. Swainson's Thrushes were only slightly selective and did not appear to forecast. By following the simple rule of avoiding only the strongest headwinds at departure, Swainson's Thrushes could survive the 1500 km flight between Alabama and Veracruz, Mexico.

Connecting the dots: Stopover strategies of an intercontinental migratory songbird in the context of the annual cycle.

The phases of the annual cycle for migratory species are inextricably linked. Yet, less than five percent of ecological studies examine seasonal interactions. In this study, we utilized stable hydrogen isotopes to geographically link individual black-and-white warblers (Mniotilta varia) captured during spring migration with breeding destinations to understand a migrant's stopover strategy in the context of other phases of the annual cycle. We found that stopover strategy is not only a function of a bird's current energetic state, but also the distance remaining to breeding destination and a bird's time-schedule, which has previously been linked to habitat conditions experienced in the preceding phase of the annual cycle. Birds in close proximity to their breeding destination accumulate additional energy reserves prior to arrival on the breeding grounds, as reflected by higher migratory condition upon arrival, higher refueling rates measured via blood plasma metabolites, and longer stopover durations compared to birds migrating to breeding destinations farther from the stopover site. However, late birds near their breeding destination were more likely to depart on the day of arrival (i.e., transients), and among birds that stopped over at the site, the average duration of stopover was almost half the time of early conspecifics, suggesting late birds are trying to catch-up with the overall time-schedule of migration for optimal arrival time on the breeding grounds. In contrast, birds with long distances remaining to breeding destinations were more likely to depart on the day of arrival and primarily used stopover to rest before quickly resuming migration, adopting similar strategies regardless of a bird's time-schedule. Our study demonstrates that migrants adjust their en route strategies in relation to their time-schedule and distance remaining to their breeding destination, highlighting that strategies of migration should be examined in the context of other phases of the annual cycle.

Body mass and wing shape explain variability in broad-scale bird species distributions of migratory passerines along an ecological barrier during stopover.

Migrating birds are under selective pressure to complete long-distance flights quickly and efficiently. Wing morphology and body mass influence energy expenditure of flight, such that certain characteristics may confer a greater relative advantage when making long crossings over ecological barriers by modifying the flight range or speed. We explored the possibility, among light (mass <50 g) migrating passerines, that species with relatively poorer flight performance related to wing shape and/or body mass have a lower margin for error in dealing with the exigencies of a long water crossing across the Gulf of Mexico and consequently minimize their travel time or distance. We found that species-mean fat-free body mass and wing tip pointedness independently explained variability among species distributions within ~50 km from the northern coast. In both spring and autumn, lighter (i.e., slower flying) species and species with more rounded wings were concentrated nearest the coastline. Our results support the idea that morphology helps to shape broad-scale bird distributions along an ecological barrier and that migration exerts some selective force on passerine morphology. Furthermore, smaller species with less-efficient flight appear constrained to stopping over in close proximity to ecological barriers, illustrating the importance of coastal habitats for small passerine migrants.

Seasonal gene expression in a migratory songbird.

The annual migration of a bird can involve thousands of kilometres of nonstop flight, requiring accurately timed seasonal changes in physiology and behaviour. Understanding the molecular mechanisms controlling this endogenous programme can provide functional and evolutionary insights into the circannual biological clock and the potential of migratory species to adapt to changing environments. Under naturally timed photoperiod conditions, we maintained captive Swainson's thrushes (Catharus ustulatus) and performed RNA sequencing (RNA-Seq) of the ventral hypothalamus and optic chiasma to evaluate transcriptome-wide gene expression changes of individuals in migratory condition. We found that 188 genes were differentially expressed in relation to migratory state, 86% of which have not been previously linked to avian migration. Focal hub genes were identified that are candidate variables responsible for the occurrence of migration (e.g. CRABP1). Numerous genes involved in cell adhesion, proliferation and motility were differentially expressed (including RHOJ, PAK1 and TLN1), suggesting that migration-related changes are regulated by seasonal neural plasticity.

Fat, weather, and date affect migratory songbirds' departure decisions, routes, and time it takes to cross the Gulf of Mexico.

Approximately two thirds of migratory songbirds in eastern North America negotiate the Gulf of Mexico (GOM), where inclement weather coupled with no refueling or resting opportunities can be lethal. However, decisions made when navigating such features and their consequences remain largely unknown due to technological limitations of tracking small animals over large areas. We used automated radio telemetry to track three songbird species (Red-eyed Vireo, Swainson's Thrush, Wood Thrush) from coastal Alabama to the northern Yucatan Peninsula (YP) during fall migration. Detecting songbirds after crossing ∼1,000 km of open water allowed us to examine intrinsic (age, wing length, fat) and extrinsic (weather, date) variables shaping departure decisions, arrival at the YP, and crossing times. Large fat reserves and low humidity, indicative of beneficial synoptic weather patterns, favored southward departure across the Gulf. Individuals detected in the YP departed with large fat reserves and later in the fall with profitable winds, and flight durations (mean = 22.4 h) were positively related to wind profit. Age was not related to departure behavior, arrival, or travel time. However, vireos negotiated the GOM differently than thrushes, including different departure decisions, lower probability of detection in the YP, and longer crossing times. Defense of winter territories by thrushes but not vireos and species-specific foraging habits may explain the divergent migratory behaviors. Fat reserves appear extremely important to departure decisions and arrival in the YP. As habitat along the GOM is degraded, birds may be limited in their ability to acquire fat to cross the Gulf.

Patterns of testosterone in three Nearctic-Neotropical migratory songbirds during spring passage.

Preparation for breeding may overlap extensively with vernal migration in long-distance migratory songbirds. Testosterone plays a central role in mediating this transition into breeding condition by facilitating changes to physiology and behavior. While changes in testosterone levels are well studied in captive migrants, these changes are less well known in free-living birds. We examined testosterone levels in free-living Nearctic-Neotropical migrants of three species during their vernal migration. Testosterone levels increased during the migratory period in males of all three species but significantly so in only two. Testosterone levels in females remained the same throughout their migration. Our results support the extensive overlap between vernal migration and breeding preparation in male songbirds. The pattern of testosterone changes during vernal migration is far from clear in females.

El Niño-Southern Oscillation is linked to decreased energetic condition in long-distance migrants.

Predicting how migratory animals respond to changing climatic conditions requires knowledge of how climatic events affect each phase of the annual cycle and how those effects carry-over to subsequent phases. We utilized a 17-year migration dataset to examine how El Niño-Southern Oscillation climatic events in geographically different regions of the Western hemisphere carry-over to impact the stopover biology of several intercontinental migratory bird species. We found that migratory birds that over-wintered in South America experienced significantly drier environments during El Niño years, as reflected by reduced Normalized Difference Vegetation Index (NDVI) values, and arrived at stopover sites in reduced energetic condition during spring migration. During El Niño years migrants were also more likely to stopover immediately along the northern Gulf coast of the southeastern U.S. after crossing the Gulf of Mexico in small suboptimal forest patches where food resources are lower and migrant density often greater than larger more contiguous forests further inland. In contrast, NDVI values did not differ between El Niño and La Niña years in Caribbean-Central America, and we found no difference in energetic condition or use of coastal habitats for migrants en route from Caribbean-Central America wintering areas. Birds over-wintering in both regions had consistent median arrival dates along the northern Gulf coast, suggesting that there is a strong drive for birds to maintain their time program regardless of their overall condition. We provide strong evidence that not only is the stopover biology of migratory landbirds influenced by events during the previous phase of their life-cycle, but where migratory birds over-winter determines how vulnerable they are to global climatic cycles. Increased frequency and intensity of ENSO events over the coming decades, as predicted by climatic models, may disproportionately influence long-distance migrants over-wintering in South America.

Effects of landscape composition and configuration on migrating songbirds: inference from an individual-based model.

The behavior of long-distance migrants during stopover is constrained by the need to quickly and safely replenish energetic reserves. Replenishing fuel stores at stopover sites requires adjusting to unfamiliar landscapes with little to no information about the distribution of resources. Despite their critical importance to the success of songbird migration, the effects of landscape composition and configuration on fuel deposition rates (FDR [g/d]), the currency of migration, has not been tested empirically. Our objectives were to understand the effects of heterogeneous landscapes on FDR of forest-dwelling songbirds during spring migration. The results of field experiments were used to parameterize a spatially explicit, individual-based model of forest songbird movement and resulting FDR. Further field experiments were used to validate the results from the individual-based model. In simulation experiments, we altered a Gulf South landscape in a factorial design to predict the effects of future patterns under different scenarios of land use change in which the abundance of high-quality hardwood habitat and the spatial aggregation of habitat varied. Simulated FDR decreased as the amount of hardwood in the landscape decreased from 41% to 22% to 12%. Further, migrants that arrived in higher-quality habitat types gained more mass. Counter to our expectations, FDR was higher with lower spatial aggregation of habitat. Differences in refueling rates may be most influenced by whether or not an individual experiences an initial searching cost after landing in poor-quality habitat. Therefore, quickly locating habitat with sufficient food resources at each stopover may be the most important factor determining a successful migration. Our findings provide empirical evidence for the argument that hardwood forest cover is a primary determinant of the quality of a stopover site in this region. This study represents the first effort to empirically quantify FDRs based on the configuration of landscapes.

Experimental evidence for the interplay of exogenous and endogenous factors on the movement ecology of a migrating songbird.

Movement patterns during songbird migration remain poorly understood despite their expected fitness consequences in terms of survival, energetic condition and timing of migration that will carry over to subsequent phases of the annual cycle. We took an experimental approach to test hypotheses regarding the influence of habitat, energetic condition, time of season and sex on the hour-by-hour, local movement decisions of a songbird during spring stopover. To simulate arrival of nocturnal migrants at unfamiliar stopover sites, we translocated and continuously tracked migratory red-eyed vireos (Vireo olivaceus) throughout spring stopover with and without energetic reserves that were released in two replicates of three forested habitat types. Migrants moved the most upon release, during which time they selected habitat characterized by greater food abundance and higher foraging attack rates. Presumably under pressure to replenish fuel stores necessary to continue migration in a timely fashion, migrants released in poorer energetic condition moved faster and further than migrants in better condition and the same pattern was true for migrants released late in spring relative to those released earlier. However, a migrant's energetic condition had less influence on their behavior when they were in poor quality habitat. Movement did not differ between sexes. Our study illustrates the importance of quickly finding suitable habitat at each stopover site, especially for energetically constrained migrants later in the season. If an initial period prior to foraging were necessary at each stop along a migrant's journey, non-foraging periods would cumulatively result in a significant energetic and time cost to migration. However, we suggest behavior during stopover is not solely a function of underlying resource distributions but is a complex response to a combination of endogenous and exogenous factors.

Test of recrudescence hypothesis for overwintering of eastern equine encephalomyelitis virus in gray catbirds.

Eastern equine encephalitis virus (EEEV; family Togaviridae, genus Alphavirus) epizootics are infrequent, but they can lead to high mortality in infected horses and humans. Despite the importance of EEEV to human and animal health, little is known about how the virus overwinters and reinitiates transmission each spring, particularly in temperate regions where infected adult mosquitoes are unlikely to survive through the winter. One hypothesis to explain the mechanism by which this virus persists from year to year is the spring recrudescence of latent virus in avian reservoir hosts. In this study, we tested the recrudescence hypothesis with gray catbirds (Dumatella carolinensis) captured in northern Ohio (July-August 2007). Birds were experimentally infected with EEEV on 1 October 2007. In January 2008, they were then exposed to exogenous testosterone and/or extended photoperiod to initiate reactivation of latent EEEV infection. All birds became viremic with EEEV, with mean viremia of 6.0 log10 plaque-forming units/ml serum occurring at 1 d postinoculation. One male in the testosterone, long-day treatment group had EEEV viral RNA in a cloacal swab collected on 18 January 2008. Otherwise, no other catbirds exhibited reactivated infections in cloacal swabs or blood. Antibody titers fluctuated over the course of the study, with lowest titers observed in January 2008, which corresponded with the lowest mean weight of the birds. No EEEV viral RNA was detected in the blood, kidney, spleen, brain, liver, and lower intestine upon necropsy at 19 wk postinfection.

A multi-scale examination of stopover habitat use by birds.

Most of our understanding of habitat use by migrating land birds comes from studies conducted at single, small spatial scales, which may overemphasize the importance of intrinsic habitat factors, such as food availability, in shaping migrant distributions. We believe that a multi-scale approach is essential to assess the influence of factors that control en route habitat use. We determined the relative importance of eight variables, each operating at a habitat-patch, landscape, or regional spatial scale, in explaining the differential use of hardwood forests by Nearctic-Neotropical land birds during migration. We estimated bird densities through transect surveys at sites near the Mississippi coast during spring and autumn migration within landscapes with variable amounts of hardwood forest cover. At a regional scale, migrant density increased with proximity to the coast, which was of moderate importance in explaining bird densities, probably due to constraints imposed on migrants when negotiating the Gulf of Mexico. The amount of hardwood forest cover at a landscape scale was positively correlated with arthropod abundance and had the greatest importance in explaining densities of all migrants, as a group, during spring, and of insectivorous migrants during autumn. Among landscape scales ranging from 500 m to 10 km radius, the densities of migrants were, on average, most strongly and positively related to the amount of hardwood forest cover within a 5 km radius. We suggest that hardwood forest cover at this scale may be an indicator of habitat quality that migrants use as a cue when landing at the end of a migratory flight. At the patch scale, direct measures of arthropod abundance and plant community composition were also important in explaining migrant densities, whereas habitat structure was of little importance. The relative amount of fleshy-fruited trees was positively related and was the most important variable explaining frugivorous migrant density during autumn. Although constraints extrinsic to habitat had a moderate role in explaining migrant distributions, our results are consistent with the view that food availability is the ultimate factor shaping the distributions of birds during stopover.

Blood parasites of Nearctic-Neotropical migrant passerine birds during spring trans-Gulf migration: impact on host body condition.

To test the hypothesis that migrants infected with blood parasites arrive on the northern coast of the Gulf of Mexico in poorer condition than uninfected birds, we examined 1705 migrant passerine birds representing 54 species of 11 families from 2 Gulf Coast sites for blood parasites. Three hundred and sixty (21.1%) were infected with 1 or more species of 4 genera of blood parasites. The prevalence of parasites was as follows: Haemoproteus spp. (11.7%), Plasmodium spp. (6.7%), Leucocytozoon spp. (1.3%), and Trypanosoma spp. (1.2%). Both prevalence and density of Haemoproteus spp. infection varied among species. We found no relationship of gender or age with the prevalence of Haemoproteus spp. infection or Plasmodium spp. infection, with the exception of the orchard oriole (Icterus spurius) for which older birds were more likely to be infected with Haemoproteus spp. than younger birds. We also found that scarlet tanagers and summer tanagers infected with species of Haemoproteus have lower fat scores than uninfected individuals and that rose-breasted grosbeaks and Baltimore orioles infected with Haemoproteus spp. have a smaller mean body mass than uninfected individuals. Blood parasites do seem to pose a physiological cost for Neotropical migrant passerines and may be important components of the ecology of these species.

Calibration of magnetic and celestial compass cues in migratory birds--a review of cue-conflict experiments.

Migratory birds use multiple sources of compass information for orientation, including the geomagnetic field, the sun, skylight polarization patterns and star patterns. In this paper we review the results of cue-conflict experiments designed to determine the relative importance of the different compass mechanisms, and how directional information from these compass mechanisms is integrated. We focus on cue-conflict experiments in which the magnetic field was shifted in alignment relative to natural celestial cues. Consistent with the conclusions of earlier authors, our analyses suggest that during the premigratory season, celestial information is given the greatest salience and used to recalibrate the magnetic compass by both juvenile and adult birds. Sunset polarized light patterns from the region of the sky near the horizon appear to provide the calibration reference for the magnetic compass. In contrast, during migration, a majority of experiments suggest that birds rely on the magnetic field as the primary source of compass information and use it to calibrate celestial compass cues, i.e. the relative saliency of magnetic and celestial cues is reversed. An alternative possibility, however, is suggested by several experiments in which birds exposed to a cue conflict during migration appear to have recalibrated the magnetic compass, i.e. their response is similar to that of birds exposed to cue conflicts during the premigratory season. The general pattern to emerge from these analyses is that birds exposed to the cue conflict with a view of the entire sunset sky tended to recalibrate the magnetic compass, regardless of whether the cue conflict occurred during the premigratory or migratory period. In contrast, birds exposed to the cue conflict in orientation funnels and registration cages that restricted their view of the region of sky near the horizon (as was generally the case in experiments carried out during the migratory season) did not recalibrate the magnetic compass but, instead, used the magnetic compass to calibrate the other celestial compass systems. If access to critical celestial cues, rather than the timing of exposure to the cue conflict (i.e. premigratory vs migratory), determines whether recalibration of the magnetic compass occurs, this suggests that under natural conditions there may be a single calibration reference for all of the compass systems of migratory birds that is derived from sunset (and possibly also sunrise) polarized light cues from the region of sky near the horizon. In cue-conflict experiments carried out during the migratory season, there was also an interesting asymmetry in the birds' response to magnetic fields shifted clockwise and counterclockwise relative to celestial cues. We discuss two possible explanations for these differences: (1) lateral asymmetry in the role of the right and left eye in mediating light-dependent magnetic compass orientation and (2) interference from the spectral and intensity distribution of skylight at sunset with the response of the light-dependent magnetic compass.

The influence of climate on the timing and rate of spring bird migration.

Ecological processes are changing in response to climatic warming. Birds, in particular, have been documented to arrive and breed earlier in spring and this has been attributed to elevated spring temperatures. It is not clear, however, how long-distance migratory birds that overwinter thousands of kilometers to the south in the tropics cue into changes in temperature or plant phenology on northern breeding areas. We explored the relationships between the timing and rate of spring migration of long-distance migratory birds, and variables such as temperature, the North Atlantic Oscillation (NAO) and plant phenology, using mist net capture data from three ringing stations in North America over a 40-year period. Mean April/May temperatures in eastern North America varied over a 5 degrees C range, but with no significant trend during this period. Similarly, we found few significant trends toward earlier median capture dates of birds. Median capture dates were not related to the NAO, but were inversely correlated to spring temperatures for almost all species. For every 1 degrees C increase in spring temperature, median capture dates of migratory birds averaged, across species, one day earlier. Lilac (Syringa vulgaris) budburst, however, averaged 3 days earlier for every 1 degrees C increase in spring temperature, suggesting that the impact of temperature on plant phenology is three times greater than on bird phenology. To address whether migratory birds adjust their rate of northward migration to changes in temperature, we compared median capture dates for 15 species between a ringing station on the Gulf Coast of Louisiana in the southern USA with two stations approximately 2,500 km to the north. The interval between median capture dates in Louisiana and at the other two ringing stations was inversely correlated with temperature, with an average interval of 22 days, that decreased by 0.8 days per 1 degrees C increase in temperature. Our results suggest that, although the onset of migration may be determined endogenously, the timing of migration is flexible and can be adjusted in response to variation in weather and/or phenology along migration routes.

Arrival fat and reproductive performance in a long-distance passerine migrant.

Long-distance passerine migrants deposit substantial fat stores to fuel their migratory journey. Many of those migratory birds arrive at their northerly breeding grounds with larger fat stores than were necessary to reach their breeding area. Both male and female American Redstarts ( Setophaga ruticilla) arrived to breed in Michigan's Upper Peninsula with fat, and females arrived with more fat than males in 2 out of 3 years. We test the hypothesis that migrants arriving at the breeding grounds with more body fat have higher reproductive success than birds arriving with little or no fat. Females, and to a lesser extent males, that arrive with fat experience gains in reproductive performance as evidenced by increased clutch size, egg volume, and nestling mass. The results have implications for understanding how events occurring during one phase of the annual cycle influence survival and/or reproductive performance in subsequent phases.