Quantitative and Qualitative Approaches to Identifying Migration Chronology in a Continental Migrant

The degree to which extrinsic factors influence migration chronology in North American waterfowl has not been quantified, particularly for dabbling ducks. Previous studies have examined waterfowl migration using various methods, however, quantitative approaches to define avian migration chronology over broad spatio-temporal scales are limited, and the implications for using different approaches have not been assessed. We used movement data from 19 female adult mallards (Anas platyrhynchos) equipped with solar-powered global positioning system satellite transmitters to evaluate two individual level approaches for quantifying migration chronology. The first approach defined migration based on individual movements among geopolitical boundaries (state, provincial, international), whereas the second method modeled net displacement as a function of time using nonlinear models. Differences in migration chronologies identified by each of the approaches were examined with analysis of variance. The geopolitical method identified mean autumn migration midpoints at 15 November 2010 and 13 November 2011, whereas the net displacement method identified midpoints at 15 November 2010 and 14 November 2011. The mean midpoints for spring migration were 3 April 2011 and 20 March 2012 using the geopolitical method and 31 March 2011 and 22 March 2012 using the net displacement method. The duration, initiation date, midpoint, and termination date for both autumn and spring migration did not differ between the two individual level approaches. Although we did not detect differences in migration parameters between the different approaches, the net displacement metric offers broad potential to address questions in movement ecology for migrating species. Ultimately, an objective definition of migration chronology will allow researchers to obtain a comprehensive understanding of the extrinsic factors that drive migration at the individual and population levels. As a result, targeted conservation plans can be developed to support planning for habitat management and evaluation of long-term climate effects.     

Leapfrog migration and residents: New migratory habits in Swedish Greylag geese

Knowledge about intraspecific and individual variation in bird migration behavior is important to predict spatiotemporal distribution, patterns of phenology, breeding success, and interactions with the surrounding environment (e.g., human livelihoods). Such variation is key to adaptive, evolutionary responses, i.e., how individuals respond spatiotemporally to the environment to maximize fitness. In this study we used GPS location data from one to three full annual cycles from 76 Greylag geese (Anser anser) to test the hypothesis that geese originating at five latitudinally separated capture sites in Sweden have different migration strategies. We also assessed individual consistency in movement strategy over consecutive annual cycles. We used the scale‐independent net squared displacement modeling framework to quantify variables of autumn and spring migration for geese from each capture site: distance, timing, and duration. Our results demonstrate a positive correlation between migration distance and latitudinal origin. Geese from the northernmost site on average migrated farther south and about 15 times as far as the short‐moving or resident geese from the two southernmost sites. Movement strategies of individual geese varied considerably both within and among capture sites. Individual consistency in movement strategy from one annual cycle to the consecutive was high in geese from the northern sites moving the farthest, whereas the resident or short‐moving geese from the southernmost sites generally showed lower or no individual consistency. These changes have come about during a time span so short (i.e., ca. 35 years or 8–10 generations) that it can unlikely be explained by classical Darwinian between‐generation adaptation. Consequently, and given that young geese follow their parents during their first migration, we presume an important role of within‐family, inter‐generation change as a driver behind the large‐scale changed migration habits in Swedish Greylag geese.     

Effects of climate on fall migration phenology of monarch butterflies departing the northeastern breeding grounds in Canada

Monarch butterflies (Danaus plexippus) undergo an iconic multi-generational migration, traveling thousands of kilometers from the summer breeding grounds in southern Canada to overwintering sites in central Mexico. This migration phenomena can be affected by climate change, which may have important implications on fitness and ultimately populations status. We investigated the long-term trends in fall migration phenology of monarchs using a 25-year dataset collected along the coast of Lake Erie in Ontario, Canada. We also investigated local long-term trends in weather covariates that have the potential to influence migration phenology at this site. Patterns in standardized daily counts of monarchs were compared with local weather covariates using two methods (i.e., monthly averages and moving windows) to assess difference in outputs between analytical approaches. Our results suggest that monarch migration timing (migration midpoint, average peak, first peak, and late passage) and weather covariates have been consistent over time, in direct contrast to a similar study in Cape May, New Jersey, which showed a significant increase in both fall temperature and a 16- to 19-day shift in monarch migration timing. Furthermore, our results differed between analytical approaches. With respect to annual variability in air temperature, our monthly average analysis suggested that for each degree increase in September air temperature, late season passage would advance 4.71 days (±1.59 SE, p = .01). However, the moving window analysis suggested that this result is likely spurious and found no significant correlations between migration timing and any weather covariates. Importantly, our results caution against extrapolating the effects of climate change on the migration phenology of the monarch across study regions and the need for more long-term monitoring efforts to better understand regional drivers of variability in migration timing.     

Mountain lions on the prairie: habitat selection by recolonizing mountain lions at the edge of their range

Mountain lions (Puma concolor) have historically experienced large‐scale range contractions, but are beginning to recolonize portions of their former range. To reach potential suitable habitats in eastern North America, mountain lions need to move across the grassland and agriculture‐dominated habitats of the Great Plains, which are different from the forested areas associated with mountain lions in western North America. To inform restoration planning in this area, it is important to understand differences in mountain lion habitat selection in this “nontraditional” grassland habitat. We tracked GPS‐collared mountain lions in the Northern Great Plains of Montana and identified movement states (localized or exploratory) using behavioral change point analysis and net‐squared displacement. We compared habitat selection between the different states using step‐selection functions that included several environmental covariates. Similar to elsewhere throughout their range, across both movement states, mountain lions tended to select forested environments that were farther from human development. In contrast to more traditionally occupied mountainous regions, mountain lions in the Great Plains selected areas of lower elevations. They selected areas both near and far from water, but avoided riparian areas and selected more rugged environments when in exploratory movement states. This suggests that mountain lions in the Northern Great Plains are utilizing river corridors, particularly those with rough or broken topography during exploratory phases. To enhance future recolonization and connectivity of mountain lions to the east of our study area, we encourage managers to maintain and restore forest fragments along river corridors in the Great Plains.     

A model-driven approach to quantify migration patterns: individual, regional and yearly differences

1.?Animal migration has long intrigued scientists and wildlife managers alike, yet migratory species face increasing challenges because of habitat fragmentation, climate change and over-exploitation. Central to the understanding migratory species is the objective discrimination between migratory and nonmigratory individuals in a given population, quantifying the timing, duration and distance of migration and the ability to predict migratory movements. 2.?Here, we propose a uniform statistical framework to (i) separate migration from other movement behaviours, (ii) quantify migration parameters without the need for arbitrary cut-off criteria and (iii) test predictability across individuals, time and space. 3.?We first validated our novel approach by simulating data based on established theoretical movement patterns. We then formulated the expected shapes of squared displacement patterns as nonlinear models for a suite of movement behaviours to test the ability of our method to distinguish between migratory movement and other movement types. 4.?We then tested our approached empirically using 108 wild Global Positioning System (GPS)-collared moose Alces alces in Scandinavia as a study system because they exhibit a wide range of movement behaviours, including resident, migrating and dispersing individuals, within the same population. Applying our approach showed that 87% and 67% of our Swedish and Norwegian subpopulations, respectively, can be classified as migratory. 5.?Using nonlinear mixed effects models for all migratory individuals we showed that the distance, timing and duration of migration differed between the sexes and between years, with additional individual differences accounting for a large part of the variation in the distance of migration but not in the timing or duration. Overall, the model explained most of the variation (92%) and also had high predictive power for the same individuals over time (69%) as well as between study populations (74%). 6.?The high predictive ability of the approach suggests that it can help increase our understanding of the drivers of migration and could provide key quantitative information for understanding and managing a broad range of migratory species.     

Can long‐distance migratory birds adjust to the advancement of spring by shortening migration distance? The response of the pied flycatcher to latitudinal photoperiodic variation

Many organisms use day length as a cue for synchronizing their life cycles with seasonal changes in environmental productivity. Under rapid climate change, however, responses to day length may become maladaptive, and photo‐responsive organisms may only be able to evade increasingly unsuitable habitats if they can accommodate to a wide range of photoperiodic conditions. A previous experiment showed that the pied flycatcher, Ficedula hypoleuca, a Palaearctic‐Afrotropical migratory bird, would strongly advance the timing of spring migration and reproductive maturation if it shifted its wintering area from sub‐Saharan Africa to the Mediterranean region. However, it is unknown whether this marked response to latitudinal variation in photoperiodic conditions is continuous over the entire range of potential wintering areas, and if a shortening of migration distance would be an effective mechanism to adjust the timing of migration to rapidly changing climatic conditions. Here, we experimentally show that a moderate northward displacement of the pied flycatcher's current wintering grounds by 10° would result in a clear advancement of the termination of prenuptial moult and the initiation of spring migratory activity and gonadal growth. However, we found no further advancement under conditions simulating higher wintering latitudes, suggesting the existence of a critical photoperiodic threshold or a steep gradual response within a narrow geographical range between 10° and 20° northern latitude. Because habitat conditions in this area are deteriorating rapidly, the potential for pied flycatchers to adjust their life cycle to changing climatic conditions by shortening the migration distance may be limited in the future.     

Improving phenological forecasting models for rape stem weevil,Ceutorhynchus napi Gyll., based on long‐term multisite datasets

New threshold‐based models to predict the start of invasion by the stem‐boring pest, the rape stem weevil (Ceutorhynchus napi Gyll.) of winter oilseed rape (Brassica napus L.), were developed and compared to published models using long‐term datasets on weather and weevil phenology from experimental locations in Germany and Luxembourg. Threshold values for daily records of maximum air temperature, mean soil temperature, sunshine duration and total precipitation were adjusted to local conditions on the date of first weevil migration in spring. Mean error and the root mean squared error were used to assess model quality, where the error is defined as the number of days between predicted and observed arrival of weevils on the crop (regardless of sign). Best model results predicted first crop invasion by rape stem weevil when the thresholds of daily maximum air temperature ≥7.8°C, mean soil temperature ≥6.6°C, daily total precipitation ≤1.0 mm and sunshine duration ≥1 h were matched. This model takes into account meteorological variables likely to influence conditions at the overwintering site of the weevils in the soil, as well as variables that may limit weevil flight. Adjusted air temperature threshold values were consistently lower for Luxembourg sites than for those optimized for Germany. A simple model relating the date of first weevil invasion to accumulated daily maximum air temperature above 0°C (from 1 January) was also evaluated. This proved less suitable for forecasting crop invasion by C. napi. We suggest that phenological models using locally adjusted meteorological‐based thresholds have the potential to offer sufficiently accurate forecasts of first immigration flights by C. napi for appropriate timing of insecticide application. In addition, the developed models are suitable tools to be used in climate change impact studies.     

Migratory pathways,stopover zones and wintering destinations of Western European Nightjars Caprimulgus europaeus

Little is known about the wintering distribution of the European Nightjar Caprimulgus europaeus. We combined geolocator and GPS‐logger data from different sites in Western Europe to analyse migration routes and migration timing of this trans‐equatorial migrant. Nightjars followed a loop migration route during which they cross two ecological barriers, and converged near common stopover zones in Northern, Central and Western Africa, where they stayed for 2–3 weeks. Nightjars used the same stopover sites as several other European migrants, relying on small and discrete wintering areas within the Democratic Republic of Congo. This confirms the importance of these specific zones and highlights the vulnerability of Western European populations to habitat loss in their non‐breeding areas.     

Spatial ecology of jaguars,pumas, and ocelots: a review of the state of knowledge

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GPS‐tracking reveals non‐breeding locations and apparent molt migration of a Black‐headed Grosbeak

Black‐headed Grosbeaks (Pheucticus melanocephalus) have been observed to undergo prebasic molt during fall in the North American Monsoon region of the southwestern United States and northwestern Mexico, but it is unknown whether molt migration is pervasive across populations of the species. During the 2014 breeding season, we GPS‐tagged (where GPS is global positioning system) nine adult Black‐headed Grosbeaks in Yosemite National Park with archival GPS tags to determine specific locations where grosbeaks breeding in Yosemite spent portions of the non‐breeding season, and to assess whether those locations were consistent with molt migration. On 2 June 2015, one of these birds, a male GPS‐tagged on 19 June 2014, was recaptured with its GPS unit still attached. Data downloaded from the unit revealed that, by 20 August 2014, the bird had moved 1300 km from Yosemite National Park to Sonora, Mexico, where it remained until at least 15 October 2014. By 24 November 2014, the grosbeak had moved >1300 m from Sonora to the Michoacán‐Jalisco border region, where it remained until the last GPS‐determined location was obtained on 24 March 2015. The seasonal timing of these movements and the length of stay in Sonora are consistent with the expected behavior of a molt‐migrating bird. Remote‐sensed enhanced vegetation index (EVI) data indicated that the grosbeak arrived in the monsoon region near the area's annual peak in EVI, and then, as the index was declining sharply, departed for the Michoacán‐Jalisco region, where the index also declined during the same period, but substantially less so than in Sonora. Climate change in the coming decades is expected to delay the annual onset of the monsoon while also accelerating the initiation of arid, summer‐like conditions throughout much of western North America, possibly yielding a temporal mismatch between fall migration and the monsoon‐driven conditions that may be critical for molt‐migrating birds.     

The z-model — A proposal for spatial and temporal modeling of visual threshold perception

By considering only the modulation transfer functions of stationary, uniformly moved, and time modulated sinusoidal gratings it is possible to derive a simple model, the z-model, for the spatio-temporal frequency behaviour of one-dimensional patterns. The transmission function of this model is a band pass function of a single coordinate z, which is a quadratic form of the spatial and temporal frequencies (rotational symmetry with respect to space and time). The model is determined by only three constants. Optionally a time phase which accounts for delay and phase distortion can be added. This model can also be derived from reaction time measurements for switched on sinusoidal gratings. With this model the response of a wide variety of spatio-temporal patterns have been calculated and compared with measured threshold data. For two-dimensional patterns orientational filtering has to be added to the model leading to a further parameter. This model predicts satisfactorily the threshold modulation for a great variety of arbitrary spatio-temporal patterns. However the absolute threshold value for aperiodic transient patterns differs slightly in direction of smaller sensitivity as compared with periodic stationary patterns. This suggests that the peak detection scheme usually used in threshold detection modeling should be replaced by an integrative mechanism.     

Movement patterns of the White-tailed Sea Eagle (Haliaeetus albicilla): post-fledging behaviour,natal dispersal onset and the role of the natal environment

Exploratory movements and natal dispersal form essential processes during early life history stages of raptors, but identifying the factors shaping individual movement decisions is challenging. Global positioning system (GPS) telemetry thereby provides a promising technique to study movement patterns on adequate spatio-temporal scales. We analysed data of juvenile White-tailed Sea Eagles Haliaeetus albicilla (WTSE) in north-east Germany (n = 24) derived from GPS tracking to extensively analyse movements between fledging and emigration from the natal territory. Our goal was to determine the time point of fledging, characterize pre-emigration movements and the onset of natal dispersal while investigating the influence of the natal environment. WTSE fledged at an average age of 72 days and showed strong excursive behaviour during the post-fledging period regarding the number, distance and duration of excursions, yet with high individual variability. Excursive behaviour did not differ between sexes. On average, WTSE left the parental territory 93 days after fledging. The quantity of excursive behaviour delayed the timing of emigration and WTSE tended to postpone their emigration when foraging water was accessible within the boundaries of their parental territory. The overall results suggest that young WTSE assess the quality of the natal environment via pre-emigration movements and stay in their territory of origin for as long as internal and external conditions allow for it. Our study is one of the first to characterize post-fledging and natal dispersal movements of young WTSE to such an extent and applies modern techniques to understand related movements in relation to the natal environment. The results emphasize the urgent necessity for the extension of currently existing nest protection periods and guaranteeing sustainable management of potential breeding and foraging grounds for WTSE. Ultimately, the results are relevant for all large raptor species sensitive to human-related disturbance, as they support the increasing importance of regulations with spatio-temporal specifications for breeding populations of large raptors in densely human-inhabited areas with increasing alteration of land.     

Improvements on GPS Location Cluster Analysis for the Prediction of Large Carnivore Feeding Activities: Ground-Truth Detection Probability and Inclusion of Activity Sensor Measures

Animal space use studies using GPS collar technology are increasingly incorporating behavior based analysis of spatio-temporal data in order to expand inferences of resource use. GPS location cluster analysis is one such technique applied to large carnivores to identify the timing and location of feeding events. For logistical and financial reasons, researchers often implement predictive models for identifying these events. We present two separate improvements for predictive models that future practitioners can implement. Thus far, feeding prediction models have incorporated a small range of covariates, usually limited to spatio-temporal characteristics of the GPS data. Using GPS collared cougar (Puma concolor) we include activity sensor data as an additional covariate to increase prediction performance of feeding presence/absence. Integral to the predictive modeling of feeding events is a ground-truthing component, in which GPS location clusters are visited by human observers to confirm the presence or absence of feeding remains. Failing to account for sources of ground-truthing false-absences can bias the number of predicted feeding events to be low. Thus we account for some ground-truthing error sources directly in the model with covariates and when applying model predictions. Accounting for these errors resulted in a 10% increase in the number of clusters predicted to be feeding events. Using a double-observer design, we show that the ground-truthing false-absence rate is relatively low (4%) using a search delay of 2–60 days. Overall, we provide two separate improvements to the GPS cluster analysis techniques that can be expanded upon and implemented in future studies interested in identifying feeding behaviors of large carnivores.     

Effects of plot vegetation diversity and spatial scale on Coccinella septempunctata movement in the absence of prey

The influence that vegetation diversity and the spatial scale of that diversity exert on insect behavior has increasingly been explored in the ecological literature, but relatively few experiments have explicitly incorporated both factors in experimental treatments. We conducted a field study designed to explore the effect of both of these factors on insect movement behavior in a broccoli agroecosystem. We caught and released seven‐spotted ladybird beetles (Coccinella septempunctata L.) in plots containing different degrees of vegetation diversity at two different spatial scales in which prey had been removed. Beetle movement was recorded at timed intervals, and move lengths and turning angles were used to generate discrete path maps for each beetle. Observed mean beetle net squared displacements were compared with predicted net squared displacements, and 95% confidence intervals were generated using a bootstrap method described by Turchin (1998 ) [Quantitative Analysis of Movement: Measuring and Modeling Population Redistribution in Animals and Plants. Sinauer Associates Inc., Sunderland, MA.]. Predicted net squared displacements underestimated beetle movement in smaller plots with both low and higher vegetation diversity for the first five move lengths, whereas no significant difference between observed and predicted net squared displacement for beetles in larger plots of either level of vegetation diversity were detected. These findings highlight the need for a better understanding of how natural enemies are influenced by vegetation diversity and the spatial scale of that vegetation in agroecosystems. The implications of these results for biological control are discussed.     

Range-wide migration corridors and non-breeding areas of a northward expanding Afro-Palaearctic migrant,the European Bee-eater Merops apiaster

Across their ranges, different populations of migratory species often use separate routes to migrate between breeding and non-breeding grounds. Recent changes in climate and land-use have led to breeding range expansions in many species but it is unclear whether these populations also establish new migratory routes, non-breeding sites and migration phenology. Thus, we compared the migration patterns of European Bee-eaters Merops apiaster from two established western (n = 5) and eastern (n = 6) breeding populations in Europe, with those from a newly founded northern population (n = 19). We aimed to relate the breeding populations to the two known non-breeding clusters in Africa, and to test for similarities of migration routes and timing between the old and new populations. Western Bee-eaters used the western flyway to destinations in West Africa; the eastern birds uniformly headed south to southern African non-breeding sites, confirming a complete separation in time and space between these long-established populations. The recently founded northern population, however, also used a western corridor, but crossed the Mediterranean further east than the western population and overwintered mainly in a new non-breeding area in southern Congo/northern Angola. The migration routes and the new non-breeding range overlapped only slightly with the western, but not with the eastern, population. In contrast, migration phenology appeared to differ between the western and both the northern and the eastern populations, with tracked birds from the western population migrating 2–4 weeks earlier. The northern population thus shares some spatial traits with western Bee-eaters, but similar phenology only with eastern population. This divergence highlights the adjustments in the timing of migration to local environmental conditions in newly founded populations, and a parallel establishment of new breeding and non-breeding sites.     

Quantifying Migration Behaviour Using Net Squared Displacement Approach: Clarifications and Caveats

Estimating migration parameters of individuals and populations is vital for their conservation and management. Studies on animal movements and migration often depend upon location data from tracked animals and it is important that such data are appropriately analyzed for reliable estimates of migration and effective management of moving animals. The Net Squared Displacement (NSD) approach for modelling animal movement is being increasingly used as it can objectively quantify migration characteristics and separate different types of movements from migration. However, the ability of NSD to properly classify the movement patterns of individuals has been criticized and issues related to study design arise with respect to starting locations of the data/animals, data sampling regime and extent of movement of species. We address the issues raised over NSD using tracking data from 319 moose (Alces alces) in Sweden. Moose is an ideal species to test this approach, as it can be sedentary, nomadic, dispersing or migratory and individuals vary in their extent, timing and duration of migration. We propose a two-step process of using the NSD approach by first classifying movement modes using mean squared displacement (MSD) instead of NSD and then estimating the extent, duration and timing of migration using NSD. We show that the NSD approach is robust to the choice of starting dates except when the start date occurs during the migratory phase. We also show that the starting location of the animal has a marginal influence on the correct quantification of migration characteristics. The number of locations per day (1–48) did not significantly affect the performance of non-linear mixed effects models, which correctly distinguished migration from other movement types, however, high-resolution data had a significant negative influence on estimates for the timing of migrations. The extent of movement, however, had an effect on the classification of movements, and individuals undertaking short- distance migrations can be misclassified as other movements such as sedentary or nomadic. Our study raises important considerations for designing, analysing and interpreting movement ecology studies, and how these should be determined by the biology of the species and the ecological and conservation questions in focus.     

Australian bird phenology: a search for climate signals

Temporal and climate‐related changes in avian phenology were assessed for seven species of south‐eastern Australia using data obtained from members of the public, naturalist groups and other organizations. Despite significantly warmer temperatures (～0.02–0.03°C per year) and reduced rainfall (～1.6–8.0 mm per year) over much of south‐eastern Australia in recent decades, most species showed no corresponding trends in their timing of migration or breeding, the notable exceptions being the grey fantail (Rhipidura fuliginosa) and the flame robin (Petroica phoenicea), which migrate through Melbourne, Victoria, during autumn and spring. In many species, however, migration or breeding timing appeared to be influenced to some extent by local, rather than regional, climate conditions, particularly local daily maximum and minimum temperatures. Whether these species will noticeably change their phenology to match projected changes in climate, perhaps when a currently unknown climate threshold is crossed, or whether these species are sufficiently flexible in their foraging strategies or food sources to be able to maintain their current timing, remains to be seen.     

Holding steady: Little change in intensity or timing of bird migration over the Gulf of Mexico

Quantifying the timing and intensity of migratory movements is imperative for understanding impacts of changing landscapes and climates on migratory bird populations. Billions of birds migrate in the Western Hemisphere, but accurately estimating the population size of one migratory species, let alone hundreds, presents numerous obstacles. Here, we quantify the timing, intensity, and distribution of bird migration through one of the largest migration corridors in the Western Hemisphere, the Gulf of Mexico (the Gulf). We further assess whether there have been changes in migration timing or intensity through the Gulf. To achieve this, we integrate citizen science (eBird) observations with 21 years of weather surveillance radar data (1995–2015). We predicted no change in migration timing and a decline in migration intensity across the time series. We estimate that an average of 2.1 billion birds pass through this region each spring en route to Nearctic breeding grounds. Annually, half of these individuals pass through the region in just 18 days, between April 19 and May 7. The western region of the Gulf showed a mean rate of passage 5.4 times higher than the central and eastern regions. We did not detect an overall change in the annual numbers of migrants (2007–2015) or the annual timing of peak migration (1995–2015). However, we found that the earliest seasonal movements through the region occurred significantly earlier over time (1.6 days decade^?1). Additionally, body mass and migration distance explained the magnitude of phenological changes, with the most rapid advances occurring with an assemblage of larger‐bodied shorter‐distance migrants. Our results provide baseline information that can be used to advance our understanding of the developing implications of climate change, urbanization, and energy development for migratory bird populations in North America.     

Proximate Cues for a Short-Distance Migratory Species: an Application of Survival Analysis

ABSTRACT Investigation of bird migration has often highlighted the importance of external factors in determining timing of migration. However, little distinction has been made between short- and long-distance migrants and between local and flight birds (passage migrants) in describing migration chronology. In addition, measures of food abundance as a proximate factor influencing timing of migration are lacking in studies of migration chronology. To address the relationship between environmental variables and timing of migration, we quantified the relative importance of proximate external factors on migration chronology of local American woodcock (Scolopax minor), a short distance migrant, using event-time analysis methods (survival analysis). We captured 1,094 woodcock local to our study sites in Michigan, Minnesota, and Wisconsin (USA) during autumn 2002–2004 and documented 786 departure dates for these birds. Photoperiod appeared to provide an initial proximate cue for timing of departure. Moon phase was important in modifying timing of departure, which may serve as a navigational aid in piloting and possibly orientation. Local synoptic weather variables also contributed to timing of departure by changing the rate of departure from our study sites. We found no evidence that food availability influenced timing of woodcock departure. Our results suggest that woodcock use a conservative photoperiod-controlled strategy with proximate modifiers for timing of migration rather than relying on abundance of their primary food, earthworms. Managing harvest pressure on local birds by adjusting season lengths may be an effective management tool with consistent migration patterns from year to year based on photoperiod.