Metapopulation models for seasonally migratory animals

Metapopulation models are widely used to study species that occupy patchily distributed habitat, but are rarely applied to migratory species, because of the difficulty of identifying demographically independent subpopulations. Here, we extend metapopulation theory to describe the directed seasonal movement of migratory populations between two sets of habitat patches, breeding and non-breeding, with potentially different colonization and extinction rates between patch types. By extending the classic metapopulation model, we show that migratory metapopulations will persist if the product of the two colonization rates exceeds the product of extinction rates. Further, we develop a spatially realistic migratory metapopulation model and derive a landscape metric-the migratory metapopulation capacity-that determines persistence. This new extension to metapopulation theory introduces an important tool for the management and conservation of migratory species and may also be applicable to model the dynamics of two host-parasite systems.     

Eco-evolutionary drivers of avian migratory connectivity

Migratory connectivity, reflecting the extent by which migrants tend to maintain their reciprocal positions in seasonal ranges, can assist in the conservation and management of mobile species, yet relevant drivers remain unclear. Taking advantage of an exceptionally large (~150,000 individuals, 83 species) and more-than-a-century-long dataset of bird ringing encounters, we investigated eco-evolutionary drivers of migratory connectivity in both short- and long-distance Afro-Palearctic migratory birds. Connectivity was strongly associated with geographical proxies of migration costs and was weakly influenced by biological traits and phylogeny, suggesting the evolutionary lability of migratory behaviour. The large intraspecific variability in avian migration strategies, through which most species geographically split into distinct migratory populations, explained why most of them were significantly connected. By unravelling key determinants of migratory connectivity, our study improves knowledge about the resilience of avian migrants to ecological perturbations, providing a critical tool to inform transboundary conservation and management strategies at the population level.     

Estimating migratory connectivity of birds when re‐encounter probabilities are heterogeneous

Understanding the biology and conducting effective conservation of migratory species requires an understanding of migratory connectivity – the geographic linkages of populations between stages of the annual cycle. Unfortunately, for most species, we are lacking such information. The North American Bird Banding Laboratory (BBL) houses an extensive database of marking, recaptures and recoveries, and such data could provide migratory connectivity information for many species. To date, however, few species have been analyzed for migratory connectivity largely because heterogeneous re‐encounter probabilities make interpretation problematic. We accounted for regional variation in re‐encounter probabilities by borrowing information across species and by using effort covariates on recapture and recovery probabilities in a multistate capture–recapture and recovery model. The effort covariates were derived from recaptures and recoveries of species within the same regions. We estimated the migratory connectivity for three tern species breeding in North America and over‐wintering in the tropics, common (Sterna hirundo), roseate (Sterna dougallii), and Caspian terns (Hydroprogne caspia). For western breeding terns, model‐derived estimates of migratory connectivity differed considerably from those derived directly from the proportions of re‐encounters. Conversely, for eastern breeding terns, estimates were merely refined by the inclusion of re‐encounter probabilities. In general, eastern breeding terns were strongly connected to eastern South America, and western breeding terns were strongly linked to the more western parts of the nonbreeding range under both models. Through simulation, we found this approach is likely useful for many species in the BBL database, although precision improved with higher re‐encounter probabilities and stronger migratory connectivity. We describe an approach to deal with the inherent biases in BBL banding and re‐encounter data to demonstrate that this large dataset is a valuable source of information about the migratory connectivity of the birds of North America.     

Geolocators reveal migratory connectivity between wintering and breeding areas of Golden‐winged Warblers

The conservation of migratory songbirds is often impeded by a lack of understanding of how populations in breeding and wintering areas are geographically linked (migratory connectivity). In recent years, light‐level geolocators have improved our understanding of migratory connectivity. Such information is valuable for evaluating how conservation efforts align between the breeding and non‐breeding areas of at‐risk species, and help to more effectively prioritize the allocation of conservation funding. Golden‐winged Warblers (Vermivora chrysoptera) are imperiled migratory songbirds, but the extent to which conservation efforts in their breeding and non‐breeding areas coincide with patterns of migratory connectivity are not well known. We used light‐level geolocators to evaluate the extent to which conservation actions targeting Golden‐winged Warblers in Nicaragua and in their breeding range in North America align with patterns of migratory connectivity. We recovered six of 22 geolocators that had been deployed on male Golden‐winged Warblers at the El Jaguar Reserve during the winter of 2015–2016. All six males migrated to breeding areas in the western Great Lakes region that includes eastern Minnesota, northern Wisconsin, southwestern Ontario, and Michigan's Upper Peninsula. All six males also had similar migration routes, with spring stopovers in southern Mexico, Guatemala, and Belize, a trans‐Gulf flight, and a stopover in the region of Louisiana, Arkansas, eastern Oklahoma, and Texas. Our results, in combination with those of previous studies, demonstrate strong migratory connectivity between portions of the breeding and winter distributions of Golden‐winged Warblers currently targeted for conservation. However, additional studies are needed to improve our understanding of the stopover ecology of Golden‐winged Warblers, especially in areas where they remain for extended periods of time. Finally, patterns of migratory connectivity revealed in our study should be used in combination with existing demographic parameters for Golden‐winged Warblers in the western Great Lakes and Nicaragua to help inform full life cycle population models for this imperiled songbird.     

Limitations and mechanisms influencing the migratory performance of soaring birds

Migration is costly in terms of time, energy and safety. Optimal migration theory suggests that individual migratory birds will choose between these three costs depending on their motivation and available resources. To test hypotheses about use of migratory strategies by large soaring birds, we used GPS telemetry to track 18 adult, 13 sub‐adult and 15 juvenile Golden Eagles Aquila chrysaetos in eastern North America. Each age‐class had potentially different motivations during migration. During spring, the migratory performance (defined here as the directness of migratory flight) of adults was higher than that of any other age‐classes. Adults also departed earlier and spent less time migrating. Together, these patterns suggest that adults were primarily time‐limited and the other two age‐classes were energy‐limited. However, adults that migrated the longest distances during spring also appeared to take advantage of energy‐conservation strategies such as decreasing their compensation for wind drift. During autumn, birds of all age‐classes were primarily energy‐minimizers; they increased the length of stopovers, flew less direct routes and migrated at a slower pace than during spring. Nonetheless, birds that departed later in autumn flew more directly, indicating that time limitations may have affected their decision‐making. During both seasons, juveniles had the lowest performance, sub‐adults intermediate performance and adults the highest performance. Our results show age‐ and seasonal variation in time and energy‐minimization strategies that are not necessarily exclusive of one another. Beyond time and energy, a complex suite of factors, including weather, experience and navigation ability, influences migratory performance and decision‐making.     

Connecting breeding and wintering grounds of Neotropical migrant songbirds using stable hydrogen isotopes: a call for an isotopic atlas of migratory connectivity

There is an overdue and urgent need to establish patterns of migratory connectivity linking breeding grounds, stopover sites, and wintering grounds of migratory birds. Such information allows more effective application of conservation efforts by applying focused actions along movement trajectories at the population level. Stable isotope methods, especially those using stable hydrogen isotope abundance in feathers (δ²H_f) combined with Bayesian assignment techniques incorporating prior information such as relative abundance of breeding birds, now provide a fast and reliable means of establishing migratory connectivity, especially for Neotropical migrants that breed in North America and molt prior to fall migration. Here we demonstrate how opportunistic sampling of feathers of 30 species of wintering birds in Cuba, Venezuela, Guatemala, Puerto Rico, and Mexico, regions that have typically been poorly sampled for estimating migratory connectivity, can be assigned to breeding areas in North America through both advanced spatial assignment to probability surfaces and through simpler map lookup approaches. Incorporating relative abundance information from the North American Breeding Bird Survey in our Bayesian assignment models generally resulted in a reduction in potential assignment areas on breeding grounds. However, additional tools to constrain longitude such as DNA markers or other isotopes would be desirable for establishing breeding or molt origins of species with broad longitudinal distributions. The isotope approach could act as a rapid means of establishing basic patterns of migratory connectivity across numerous species and populations. We propose a large‐scale coordinated sampling effort on the wintering grounds to establish an isotopic atlas of migratory connectivity for North American Neotropical migrants and suggest that isotopic variance be considered as a valuable metric to quantify migratory connectivity. This initiative could then act as a strategic template to guide further efforts involving stable isotopes, light‐sensitive geolocators, and other technologies.     

Novel statistical methods for integrating genetic and stable isotope data to infer individual‐level migratory connectivity

Methods for determining patterns of migratory connectivity in animal ecology have historically been limited due to logistical challenges. Recent progress in studying migratory bird connectivity has been made using genetic and stable‐isotope markers to assign migratory individuals to their breeding grounds. Here, we present a novel Bayesian approach to jointly leverage genetic and isotopic markers and we test its utility on two migratory passerine bird species. Our approach represents a principled model‐based combination of genetic and isotope data from samples collected on the breeding grounds and is able to achieve levels of assignment accuracy that exceed those of either method alone. When applied at large scale the method can reveal specific migratory connectivity patterns. In Wilson's warblers (Wilsonia pusilla), we detect a subgroup of birds wintering in Baja that uniquely migrate preferentially from the coastal Pacific Northwest. Our approach is implemented in a way that is easily extended to accommodate additional sources of information (e.g. bi‐allelic markers, species distribution models, etc.) or adapted to other species or assignment problems.     

迁徙鸟类对中途停歇地的利用及迁徙对策

中途停歇地是迁徙鸟类在繁殖地和非繁殖地之间的联系枢纽,对于迁徙鸟类完成其完整的生活史过程具有重要作用。从鸟类的迁徙对策、中途停歇地的选择、鸟类在中途停歇地的停留时间、体重变化和种群特征以及中途停歇地的环境状况等方面,回顾了中途停歇生态学在近年来的研究进展,并提出了在迁徙对策理论的实验研究,小型鸟类在中途停歇地的停歇时间及体重变化的准确确定等目前有待解决的问题。     

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.     

BirdOriTrack: a new video‐tracking program for orientation research with migratory birds

Experimental research on the orientation of migratory songbirds is traditionally conducted using orientation funnels or automatic registration cages that record the directional activity of individual birds on paper or a computer. Most traditionally used funnel and cage designs do not permit investigators to observe detailed behavior of the birds and, therefore, we have gained little insight into the actual behavior of birds while they are exhibiting migratory restlessness and making directional choices. Such behavior can only be studied by direct observation or by video filming. Here, we present BirdOriTrack, a video‐tracking program for extracting time‐resolved, positional data of birds (and potentially other animal species) to determine their orientation relative to the center of a circular cage/funnel. With relatively inexpensive cameras, recording equipment, and cages, orientation experiments can easily be conducted and analyzed using BirdOriTrack. The program is designed to be flexible, allowing analysis of orientation behavior of birds of any size in different cage designs and in both controlled laboratory settings and field‐based studies. To demonstrate the program's utility, we show the results of preliminary field experiments on several species of migratory birds captured at a migration monitoring station. BirdOriTrack is freely available at http://canmove.lu.se/birdoritrack .     

Feather deuterium measurements reveal origins of migratory western loggerhead shrikes (Lanius ludovicianus excubitorides) wintering in Mexico

Understanding the winter distributions of migrant birds is important because productivity and recruitment are influenced by conditions at several locations and periods in the life cycle of individuals. The western loggerhead shrike, Lanius ludovicianus excubitorides , is a threatened species in Canada, and its decline is attributed to potential limitations on the wintering grounds. We examined patterns of stable-hydrogen isotope (δD) distributions in feathers of loggerhead shrikes, primarily of L. l. excubitorides , during winter at three regions in north and central Mexico, to establish relative abundance and origins of migrants. We also investigated potential movements of Mexican winter resident individuals. Using shrike museum specimens of known summer provenance, a shrike deuterium base map for Mexico was developed from isotopic measurement of feathers of resident shrikes and use of a recently established feather base map for raptors in North America. Stable hydrogen isotope analyses of inner secondary feather (s9) of all loggerhead shrikes examined in Mexico during winter indicated that north-central (Region A), north-eastern (Region B) and south-central (Region C) sites in Mexico consisted of 28.1%, 73.7% and 63.8% of migrant individuals from northern breeding grounds, respectively. Isotopic evidence suggested movements of a few local residents birds (7.9%) into the Chihuahuan desert from south-western USA and north-eastern Mexico to winter.     

Accipiter hawks of the Laurentian Upland and the Interior Plains undertake the longest migrations: insights from birds banded or recovered in Veracruz,Mexico

Bird banding has allowed us to understand diverse aspects of the life histories of migratory raptors. However, most banding stations are located at northern latitudes so what we know about the movements of these raptors is biased toward higher latitudes, primarily from Canada and the United States, leaving important gaps in our knowledge of their movements at lower latitudes. Our objective was to describe the migratory movements of Sharp-shinned (Accipiter striatus) and Cooper’s (A. cooperii) hawks based on banding and recapture records of birds that migrate through Veracruz, Mexico. More specifically, we sought to determine their breeding, migration, and non-breeding locations, estimate their migration distances, and contribute to a better understanding of their migration patterns. With a total of 80 records, we calculated migration distances and used Kernel Density Estimation analyses to identify where these hawks were recaptured or recovered by season. The distribution of recaptures and recoveries largely coincided with breeding locations in the Laurentian Upland and the Interior Plains physiographic regions. All migration records follow a trajectory that extends from the midwestern United States to the Gulf coastal plain of Mexico. The mean breeding season migration distance to Veracruz was 3374 km (a difference of 27 degrees of latitude) for Sharp-shinned Hawks and 2926 km (a difference of 25 degrees of latitude) for Cooper’s Hawks. Our non-breeding records indicate that populations of Accipiter hawks from these North American populations migrate the longest distances to reach Central America, the southernmost distribution of their migratory populations. Distances covered by both species represent round-trip migrations that may be as long as 10,000 km. Our results support those of previous studies and illustrate how continental physiography influences the migration routes, migratory behavior, and migratory connectivity of these hawks.     

Regional climate on the breeding grounds predicts variation in the natal origin of monarch butterflies overwintering in Mexico over 38 years

Addressing population declines of migratory insects requires linking populations across different portions of the annual cycle and understanding the effects of variation in weather and climate on productivity, recruitment, and patterns of long‐distance movement. We used stable H and C isotopes and geospatial modeling to estimate the natal origin of monarch butterflies (Danaus plexippus) in eastern North America using over 1000 monarchs collected over almost four decades at Mexican overwintering colonies. Multinomial regression was used to ascertain which climate‐related factors best‐predicted temporal variation in natal origin across six breeding regions. The region producing the largest proportion of overwintering monarchs was the US Midwest (mean annual proportion = 0.38; 95% CI: 0.36–0.41) followed by the north‐central (0.17; 0.14–0.18), northeast (0.15; 0.11–0.16), northwest (0.12; 0.12–0.16), southwest (0.11; 0.08–0.12), and southeast (0.08; 0.07–0.11) regions. There was no evidence of directional shifts in the relative contributions of different natal regions over time, which suggests these regions are comprising the same relative proportion of the overwintering population in recent years as in the mid‐1970s. Instead, interannual variation in the proportion of monarchs from each region covaried with climate, as measured by the Southern Oscillation Index and regional‐specific daily maximum temperature and precipitation, which together likely dictate larval development rates and food plant condition. Our results provide the first robust long‐term analysis of predictors of the natal origins of monarchs overwintering in Mexico. Conservation efforts on the breeding grounds focused on the Midwest region will likely have the greatest benefit to eastern North American migratory monarchs, but the population will likely remain sensitive to regional and stochastic weather patterns.     

Parasites hinder monarch butterfly flight: implications for disease spread in migratory hosts

Monarch butterflies (Danaus plexippus) are parasitized by the protozoan Ophryocystis elektroscirrha throughout their geographical range. Monarchs inhabiting seasonally fluctuating environments migrate annually, and parasite prevalence is lower among migratory relative to non‐migratory populations. One explanation for this pattern is that long‐distance migration weeds out infected animals, thus reducing parasite prevalence and transmission between generations. In this study we experimentally infected monarchs from a migratory population and recorded their long‐distance flight performance using a tethered flight mill. Results showed that parasitized butterflies exhibited shorter flight distances, slower flight speeds, and lost proportionately more body mass per km flown. Differences between parasitized and unparasitized monarchs were generally not explained by individual variation in wing size, shape, or wing loading, suggesting that poorer flight performance among parasitized hosts was not directly caused by morphological constraints. Effects of parasite infection on powered flight support a role for long‐distance migration in dramatically reducing parasite prevalence in this and other host–pathogen systems.