Pre-migratory flights in migrant songbirds: the ecological and evolutionary importance of understudied exploratory movements.

Across the animal kingdom, from honeybees to cranes to beavers, exploratory movements to exploit resources, scout prospective territories, or otherwise gain valuable experiences and information that promote fitness have been documented. For example, exploratory movements to investigate potential dispersal targets have been observed in roe deer, Northern cardinals, and tigers alike. However, despite how widespread these movements are, a cohesive definition of exploratory movements has been lacking. We first provide a clear definition of exploratory movements, and use one particular group-migratory songbirds-to catalogue exploratory movements across the annual cycle. The exceptional mobility of migratory songbirds results in exploratory movements not only at a local scale, but also on a regional scale, both in and out of the breeding season. We review the extent to which these movements are made within this group, paying particular attention to how such movements confer fitness benefits, as by securing high-quality territories, prospecting for extra-pair paternity, or even exploiting ephemeral resources. We then zoom in one step further to a particular exploratory movement that has been, to date, almost completely overlooked within this group: that of pre-migratory flights. These flights, which occur during the transitional period between the stationary breeding period and the onset of migration, occur at night and may not be made by all individuals in a population-reasons why these flights have been heretofore critically understudied. We provide the first definition for this behaviour, summarise the current knowledge of this cryptic movement, and hypothesise what evolutionary/ecological advantages conducting it may confer to the individuals that undertake it. As these flights provide experience to the individuals that undertake them, we expect that birds that make pre-migratory flights are better equipped to survive migration (direct fitness benefits) and, due to orientation/navigation abilities, may also reach preferred territories on breeding and wintering grounds faster (indirect fitness benefits). We hope to encourage ecologists to consider such hidden movements in their research concepts and to enhance the framework of movement ecology by this behaviour due to its presumed high biological importance to the annual cycle of birds.

Songbirds initiate migratory flights synchronously relative to civil dusk.

BACKGROUND: Each spring and fall billions of songbirds depart on nocturnal migrations across the globe. Theory suggests that songbirds should depart on migration shortly after sunset to maximize their potential for nightly flight duration or to time departure with the emergence of celestial cues needed for orientation and navigation. Although captive studies have found that songbirds depart during a narrow window of time after sunset, observational studies have found that wild birds depart later and more asynchronously relative to sunset than predicted. METHODS: We used coded radio tags and automated radio-telemetry to estimate the time that nearly 400 individuals from nine songbird species departed their breeding or wintering grounds across North America. We also assessed whether each species was most likely beginning long-distance migratory flights at departure or instead first making non-migratory regional flights. We then explored variation in nocturnal departure time by post-departure movement type, species, age, sex, and season. RESULTS: We found that 90% of individuals from species that were likely initiating long-distance migratory flights departed within 69 min of civil dusk, regardless of species, season, age, or sex. By contrast, species that likely first made non-migratory regional movements away from the migratory destination departed later and more asynchronously throughout the night. Regardless of post-departure movement type, 98% of individuals departed after civil dusk but otherwise showed no preference in relation to twilight phase. CONCLUSIONS: Although the presence of celestial orientation cues at civil dusk may set a starting point for departure each night, the fact that species likely beginning long-distance migration departed earlier and more synchronously relative to civil dusk than those first making non-migratory regional movements is consistent with the hypothesis that departing promptly after civil dusk functions to maximize the potential for nightly flight duration and distance. By studying the onset of migration, our study provides baseline information about departure decisions that may enhance our understanding of departure timing throughout migration.

Atmospheric pressure predicts probability of departure for migratory songbirds.

BACKGROUND: Weather can have both delayed and immediate impacts on animal populations, and species have evolved behavioral adaptions to respond to weather conditions. Weather has long been hypothesized to affect the timing and intensity of avian migration, and radar studies have demonstrated strong correlations between weather and broad-scale migration patterns. How weather affects individual decisions about the initiation of migratory flights, particularly at the beginning of migration, remains uncertain. METHODS: Here, we combine automated radio telemetry data from four species of songbirds collected at five breeding and wintering sites in North America with hourly weather data from a global weather model. We use these data to determine how wind profit, atmospheric pressure, precipitation, and cloud cover affect probability of departure from breeding and wintering sites. RESULTS: We found that the probability of departure was related to changes in atmospheric pressure, almost completely regardless of species, season, or location. Individuals were more likely to depart on nights when atmospheric pressure had been rising over the past 24 h, which is predictive of fair weather over the next several days. By contrast, wind profit, precipitation, and cloud cover were each only informative predictors of departure probability in a single species. CONCLUSIONS: Our results suggest that individual birds actively use weather information to inform decision-making regarding the initiation of departure from the breeding and wintering grounds. We propose that birds likely choose which date to depart on migration in a hierarchical fashion with weather not influencing decision-making until after the departure window has already been narrowed down by other ultimate and proximate factors.

Spatial structure in migration routes maintained despite regional convergence among eastern populations of Swainson's Thrushes.

BACKGROUND: Migratory connectivity links the different populations across the full cycle and across the species range and may lead to differences in survival among populations. Studies on spatial and temporal migratory connectivity along migration routes are rare, especially for small migratory animals. METHODS: We used an automated radio-telemetry array to assess migratory connectivity en route and between early and later stages of the fall migration of the eastern populations of Swainson's Thrush, and to assess the variation of migration pace between consecutive detection from the different receiving stations along the migratory journey. We tracked 241 individuals from across eastern Canada to determine if populations were mixing around the Gulf of Mexico. We also tested the influence of tagging longitude, latitude and age on migration pace. RESULTS: Migration routes varied and converged towards the northeast coast of the Gulf of Mexico, but in this region, populations maintained finer-scale spatial structure. Migration pace increased as birds progressed south, independent of age and tagging site. CONCLUSIONS: We showed that for songbirds, migratory connectivity can be maintained at fine spatial scales despite the regional convergence of populations, highlighting the importance of detailed spatial tracking for identification of population specific migration routes. Overall, our study provides a portrait of migratory movements of eastern Swainson's Thrush and a framework for understanding spatial structure in migration routes for other species.

Nocturnal departure timing in songbirds facing distinct migratory challenges.

Most migratory songbirds travel between their breeding areas and wintering grounds through a series of nocturnal flights. The timing of their departures defines the potential flight duration and thus the distance covered during a migratory night. Yet, migratory songbirds show substantial variation in their nocturnal departure timing. With this study, we aim to assess whether the respective challenges of the migration route, namely its distance and nature, help to explain this variation. At a stopover site, we caught Northern Wheatears (Oenanthe oenanthe) of two subspecies that differ in distance and nature of their onward migration route in spring, but not in autumn. We determined the start of their nocturnal migratory restlessness during short-term captivity, and radiotracked their nocturnal departure timing after release in both migration seasons. Northern Wheatears started their nocturnal migratory restlessness earlier when facing a long remaining migration distance and an extended sea barrier in spring. Individual departure directions generally affected the nocturnal departure timing with early departures being directed towards the respective migratory destination. In spring, this pattern was predominantly found in birds carrying relatively large fuel stores, but was absent in lean birds. At the same time, birds facing a short remaining migration distance and no extended sea barrier strongly reacted to relatively large fuel stores by an early start of nocturnal migratory behaviour (migratory restlessness and departure timing), whereas this reaction was not found in birds facing a long remaining migration distance and sea barrier. These results suggest that the basic diel schedule of birds' migratory activity is adapted to the onward migration route. Further, they suggest that birds adjust their behavioural response, that is start of nocturnal migratory behaviour, to fuel stores in relation to their impending migratory challenges. This is a substantial step in understanding variation of nocturnal departure timing and its adjustments in migratory songbirds. Further, it emphasizes the importance of interpreting birds' nocturnal migratory behaviour in the respective ecological context.

Migratory blackpoll warblers (Setophaga striata) make regional-scale movements that are not oriented toward their migratory goal during fall.

BACKGROUND: Regional scale movement patterns of songbirds are poorly known largely due to difficulties tracking small organisms at broad scales. Using an array of over 100 automated radio telemetry towers, we followed Blackpoll Warblers (Setophaga striata) during fall migration in the Gulf of Maine region, and assessed how their regional scale movement pathways varied with age, distance to natal origin, and capture date. RESULTS: Many individuals had movement paths that were not oriented towards their migratory goal ('indirect movement patterns'), regardless of age, distance to natal origin, or time of season. The probability of moving in indirect patterns, and the total tracking duration, decreased with capture date. The extent of indirect movement patterns varied considerably between individuals. Excluding direct flight patterns consistent with traditional migratory movements, adults tended to make more flights and moved in more tortuous patterns than hatch-years. Adults and individuals from more westerly natal origins were more likely to move south-west through time. CONCLUSIONS: A greater proportion of individuals made movements that were not oriented towards the migratory than expected. A decrease in tracking duration with capture date indicates that individuals prioritize time as the season progresses. The shorter, indirect movement patterns may be a more complete representation of 'reverse migration' at a barrier or 'landscape-scale stopovers movements'. The longer distances travelled are inconsistent with expected behaviour, even in front of a barrier. The extent of movement we observed indirectly suggests that flight is not as costly to individuals in a migratory state as is commonly assumed. Since adults were observed to move more than hatch-years, we suggest that the indirect movement patterns we observed are not accidental, and may provide some advantage to the individuals that undertake them.

Fuel loads acquired at a stopover site influence the pace of intercontinental migration in a boreal songbird.

Long-distance migratory organisms are under strong selection to migrate quickly. Stopovers demand more time than flying and are used by individuals to refuel during migration, but the effect of fuel loads (fat) acquired at stopover sites on the subsequent pace of migration has not been quantified. We studied stopover behaviour of Grey-cheeked Thrush (Catharus minimus) at a site in northern Colombia and then tracked their migration using an intercontinental radio-telemetry array. Tracking confirmed long-distance flights of more than 3000 km, highlighting the key importance of a single stopover site to the migration strategy of this species. Our results suggest that these songbirds behave as time-minimizers as predicted by optimal migration theory, and that fuel loads acquired at this South American stopover site, together with departure date, carry-over to influence the pace of migration, contributing to differences in travel time of up to 30 days in birds subsequently detected in the U. S. and Canada. Such variation in the pace of migration arising from a single stopover site, likely has important fitness consequences and suggests that identifying important fuelling sites will be essential to effectively conserve migratory species.

Migratory orientation in a narrow avian hybrid zone.

BACKGROUND: Zones of contact between closely related taxa with divergent migratory routes, termed migratory divides, have been suggested as areas where hybrid offspring may have intermediate and inferior migratory routes, resulting in low fitness of hybrids and thereby promoting speciation. In the Rocky Mountains of Canada there is a narrow hybrid zone between Audubon's and myrtle warblers that is likely maintained by selection against hybrids. Band recoveries and isotopic studies indicate that this hybrid zone broadly corresponds to the location of a possible migratory divide, with Audubon's warblers migrating south-southwest and myrtle warblers migrating southeast. We tested a key prediction of the migratory divide hypothesis: that genetic background would be predictive of migratory orientation among warblers in the center of the hybrid zone. METHODS: We recorded fall migratory orientation of wild-caught migrating warblers in the center of the hybrid zone as measured by video-based monitoring of migratory restlessness in circular orientation chambers. We then tested whether there was a relationship between migratory orientation and genetic background, as measured using a set of species-specific diagnostic genetic markers. RESULTS: We did not detect a significant association between orientation and genetic background. There was large variation among individuals in orientation direction. Mean orientation was towards the NE, surprising for birds on fall migration, but aligned with the mountain valley in which the study took place. CONCLUSIONS: Only one other study has directly analyzed migratory orientation among naturally-produced hybrids in a migratory divide. While the other study showed an association between genetic background and orientation, we did not observe such an association in yellow-rumped warblers. We discuss possible reasons, including the possibility of a lack of a strong migratory divide in this hybrid zone and/or methodological limitations that may have prevented accurate measurements of long-distance migratory orientation.

Corticosterone and timing of migratory departure in a songbird.

Bird migration entails replenishing fuel stores at stopover sites. There, individuals make daily decisions whether to resume migration, and must also decide their time of departure. Variation in departure timing affects the total time required to complete a migratory journey, which in turn affects fitness through arrival time at the breeding and wintering grounds. It is well established that stopover departure decisions are based on cues from innate rhythms, intrinsic factors and extrinsic factors. Yet, virtually nothing is known about the physiological mechanism(s) linking these cues to departure decisions. Here, we show for a nocturnal migratory songbird, the northern wheatear (Oenanthe oenanthe), that baseline corticosterone levels of birds at stopover increased both over the migratory season and with wind assistance towards the migratory destination. Corticosterone in turn predicted departure probability; individuals with high baseline corticosterone levels were more likely to resume migration on a given night. Corticosterone further predicted the departure time within the night, with high baseline levels being associated with early departures. These novel findings indicate that corticosterone may be mediating between departure cues and the timing of departure from a stopover site, which is a major step towards understanding the hormonal control of animal migration.

Towards a conceptual framework for explaining variation in nocturnal departure time of songbird migrants.

Most songbird migrants travel between their breeding areas and wintering grounds by a series of nocturnal flights. The exact nocturnal departure time for these flights varies considerably between individuals even of the same species. Although the basic circannual and circadian rhythms of songbirds, their adaptation to migration, and the factors influencing the birds' day-to-day departure decision are reasonably well studied, we do not understand how birds time their departures within the night. These decisions are crucial, because the nocturnal departure time defines the potential flight duration of the migratory night. The distances covered during the nocturnal migratory flights in the course of migration in turn directly affect the overall speed of migration. To understand the factors influencing the arrival of the birds in the breeding/wintering areas, we need to investigate the mechanisms that control nocturnal departure time. Here, we provide the first conceptual framework for explaining the variation commonly observed in this migratory trait. The basic schedule of nocturnal departure is likely regulated by both the circannual and circadian rhythms of the innate migration program. We postulate that the endogenously controlled schedule of nocturnal departures is modified by intrinsic and extrinsic factors. So far there is only correlative evidence that birds with a high fuel load or a considerable increase in fuel load and significant wind (flow) assistance towards their migratory goal depart early within the night. In contrast, birds migrating with little fuel and under unfavorable wind conditions show high variation in their nocturnal departure time. The latter may contain an unknown proportion of nocturnal movements not directly related to migratory flights. Excluding such movements is crucial to clearly identify the main drivers of the variation in nocturnal departure time. In general we assume that the observed variation in the nocturnal departure time is explained by individually different reactions norms of the innate migration program to both intrinsic and extrinsic factors.

An Investigation Into the Integrity of Fit of Provisional Crowns Using Current Proprietary Temporary Crown Materials.

Three methods of direct provisional crown construction were investigated for accuracy of marginal fit. A modified proprietary crown coping was compared to Bis GMA and isobutyl methacrylate resin provisional crowns with margins modified by using a flowable composite and 'bead on' isobutyl methacrylate respectively. Measurement was at 50x magnification at seven sites over the fit surface. Data was analyzed using SPSS version 13.0.1 and measurement compared using the Mann Whitney test set at a significance level of 0.05. Reliability was checked using the Bland Altman test. Statistical significant differences were found between the three groups. The order of best fit was Bis-GMA and flowable composite > isobutyl methacrylate with 'bead on' margins > Bis-GMA modified implant temporary coping. The clinical significance is that the Bis GMA and flowable composite combination can be used with equal confidence to traditional methods of temporarisation.

Prosthodontic Rehabilitation for a Patient with Ellis-Van Creveld Syndrome: A Case Report.

Patients suffering from Ellis-Van-Creveld syndrome are a challenge for dental management. Aesthetics are a major concern with limited manual dexterity, making choice of treatment critical. A 38 year old female diagnosed with Ellis-Van-Creveld syndrome presented with stained teeth and un-aesthetic smile and related that to her low self-esteem and depression. Intra-oral examination revealed mal-aligned megadont central incisors in the maxillary arch, fused mandibular canines with laterals and missing central incisors with space discrepancy and pronounced reverse over jet and overbite. Treatment involved non-surgical periodontal management, fabrication of veneers and dentine bonded crowns for maxillary anteriors and fixed-fixed resin retained bridge for mandibular arch.

Differential fall migratory routes of adult and juvenile Ipswich Sparrows (Passerculus sandwichensis princeps).

BACKGROUND: Island breeding birds present an ideal system for studying migratory movements in passerines because their populations are clearly demarcated, and individuals must depart on migration from a single location. The Ipswich Sparrow (Paserculus sandwichensis princeps) is a subspecies of the Savannah Sparrow that breeds exclusively on Sable Island, Nova Scotia, Canada and winters along the Atlantic coast of North America. We used a network of 34 automated VHF telemetry receivers to track radio-tagged adult and juvenile Ipswich Sparrows from their breeding island southward through the first half of their fall migratory journey. RESULTS: We compared adult to juvenile timing and routes. We show that juveniles leave the island approximately 24 days prior to adults and remain temporally separated from them during migration through Nova Scotia. Juveniles have different overwater orientations that result in migratory routes with shorter ocean crossings and a longer overall distance travelled compared to adults. Juveniles also have more frequent and longer stopovers, and displayed some reverse migration. CONCLUSION: We demonstrate that migratory routes differ between adults and juveniles, suggesting that routes change as individuals age, possibly through learning or social interactions. These differential routes also suggest that sparrows experience risk in different ways with juveniles selecting shorter overwater flights with less navigational risk at the cost of increased time spent in migration.

Adult and hatch-year blackpoll warblers exhibit radically different regional-scale movements during post-fledging dispersal.

Using a broad-scale automated telemetry array, we explored post-fledging movements of blackpoll warblers breeding in Atlantic Canada. We sought to determine the full spatial scale of post-fledging dispersal, to assess support for three hypotheses for regional-scale post-fledging movement, and to determine whether learning influenced movement during this period. We demonstrated that both young and adults moved over distances more than 200 km prior to initiating migration. Adults moved southwest, crossing the Gulf of Maine (GOM), consistent with the commencement of migration hypothesis. Hatch-year birds exhibited less directional movements constrained geographically by the GOM. Their movements were most consistent with exploration hypotheses--that young birds develop a regional-scale map to aid in habitat selection, natal dispersal and subsequent migrations.

An Investigation into the Accuracy of Two Currently Available Dental Impression Materials in the Construction of Cobalt-Chromium Frameworks for Removable Partial Dentures.

This study investigated the suitability of irreversible hydrocolloid as an impression material for cobalt-chromium framework construction. Scans of casts derived from (1) alginate and (2) addition-cured polyvinylsiloxane impressions were superposed on to a control. The differences within and between groups were compared at fixed landmarks. The investigation revealed a high degree of scan coincidence within and between groups. However, certain features, such as undercuts, resulted in a lower degree of scan coincidence. Irreversible hydrocolloid appears to be a viable alternative to addition-cured polyvinyl-siloxane as an impression material for cobalt-chromium framework construction.

Automated telemetry reveals age specific differences in flight duration and speed are driven by wind conditions in a migratory songbird.

BACKGROUND: Given that winds encountered on migration could theoretically double or half the energy expenditure of aerial migrants, there should be strong selection on behaviour in relation to wind conditions aloft. However, evidence suggests that juvenile songbirds are less choosy about wind conditions at departure relative to adults, potentially increasing energy expenditure during flight. To date, there has yet to be a direct comparison of flight efficiency between free-living adult and juvenile songbirds during migration in relation to wind conditions aloft, likely because of the challenges of following known aged individual songbirds during flight. We used an automated digital telemetry array to compare the flight efficiency of adult and juvenile Savannah sparrows (Passerculus sandwichensis) as they flew nearly 100 km during two successive stages of their fall migration; a departure flight from their breeding grounds out over the ocean and then a migratory flight along a coast. Using a multilevel path modelling framework, we evaluated the effects of age, flight stage, tailwind component, and crosswind component on flight duration and groundspeed. RESULTS: We found that juveniles departed under wind conditions that were less supportive relative to adults and that this resulted in juveniles taking 1.4 times longer to complete the same flight trajectories as adults. We did not find an effect of age on flight duration or groundspeed after controlling for wind conditions aloft, suggesting that both age groups were flying at similar airspeeds. We also found that groundspeeds were 1.7 times faster along the coast than over the ocean given more favourable tailwinds along the coast and because birds appeared to be climbing in altitude over the ocean, diverting some energy from horizontal to vertical movement. CONCLUSIONS: Our results provide the first evidence that adult songbirds have considerably more efficient migratory flights than juveniles, and that this efficiency is driven by the selection of more supportive tailwind conditions aloft. We suggest that the tendency for juveniles to be less choosy about wind conditions at departure relative to adults could be adaptive if the benefits of having a more flexible departure schedule exceed the time and energy savings realized during flight with more supportive winds.

Modeling Systematic Change in Stopover Duration Does Not Improve Bias in Trends Estimated from Migration Counts.

The use of counts of unmarked migrating animals to monitor long term population trends assumes independence of daily counts and a constant rate of detection. However, migratory stopovers often last days or weeks, violating the assumption of count independence. Further, a systematic change in stopover duration will result in a change in the probability of detecting individuals once, but also in the probability of detecting individuals on more than one sampling occasion. We tested how variation in stopover duration influenced accuracy and precision of population trends by simulating migration count data with known constant rate of population change and by allowing daily probability of survival (an index of stopover duration) to remain constant, or to vary randomly, cyclically, or increase linearly over time by various levels. Using simulated datasets with a systematic increase in stopover duration, we also tested whether any resulting bias in population trend could be reduced by modeling the underlying source of variation in detection, or by subsampling data to every three or five days to reduce the incidence of recounting. Mean bias in population trend did not differ significantly from zero when stopover duration remained constant or varied randomly over time, but bias and the detection of false trends increased significantly with a systematic increase in stopover duration. Importantly, an increase in stopover duration over time resulted in a compounding effect on counts due to the increased probability of detection and of recounting on subsequent sampling occasions. Under this scenario, bias in population trend could not be modeled using a covariate for stopover duration alone. Rather, to improve inference drawn about long term population change using counts of unmarked migrants, analyses must include a covariate for stopover duration, as well as incorporate sampling modifications (e.g., subsampling) to reduce the probability that individuals will be detected on more than one occasion.

Transoceanic migration by a 12 g songbird.

Many fundamental aspects of migration remain a mystery, largely due to our inability to follow small animals over vast spatial areas. For more than 50 years, it has been hypothesized that, during autumn migration, blackpoll warblers (Setophaga striata) depart northeastern North America and undertake a non-stop flight over the Atlantic Ocean to either the Greater Antilles or the northeastern coast of South America. Using miniaturized light-level geolocators, we provide the first irrefutable evidence that the blackpoll warbler, a 12 g boreal forest songbird, completes an autumn transoceanic migration ranging from 2270 to 2770 km (mean ± s.d.: 2540 ± 257) and requiring up to 3 days (62 h ± 10) of non-stop flight. This is one of the longest non-stop overwater flights recorded for a songbird and confirms what has long been believed to be one of the most extraordinary migratory feats on the planet.