Fall Migration and Habitat Use of American Woodcock in the Central United States

Abstract: Little is known about the migration ecology of the American woodcock (Scolopax minor). From 2001 to 2003, we began a 3-year study to document woodcock fall migration routes, rates, and habitat use from Minnesota, Wisconsin, and Michigan, USA. Some 586 radiomarked woodcock initiated migration. During 224 hours of aerial telemetry, we located 42 radiomarked woodcock in 6 states. Using locations of radiomarked birds, we speculated woodcock migration routes in the central United States. Stopover duration often exceeded 4 days, with some birds stopping longer than a week. Radiomarked birds were located in upland habitats more frequently than bottomland habitats, and used a higher proportion of mature forest than expected. A Geographic Information System was used to map potential woodcock habitat in the Central Region. Based on our results, we identified possible fall migration routes and priority areas for woodcock management in the Central Region. Our results should be used by land managers to prioritize future land acquisition and management of woodcock habitat.     

Sex-specific differences and long-term trends in habitat selection of American woodcock

Effective wildlife management requires an understanding of how individuals select environmental factors, although few studies assess how habitat selection may differ over time or between sexes. During the post-breeding period (15 May to 1 Sep), we tracked 146 male American woodcock (Scolopax minor) in Rhode Island, USA, from 2010–2021 to assess how habitat selection varied over time, and 17 females and 51 males during the final 2 years of the study to document sex-specific differences in habitat selection. Males generally had smaller home ranges (35.0 ± 10.7 [margin of error] ha) and preferred habitat mosaics that consisted of forested wetlands, young forest patches, areas of deciduous forest, moist soils with gentle slopes, and riparian corridors. We detected subtle differences between sexes in selection for wetland young forest, upland young forest, percent slope, distance to upland young forest, distance to streams, and distance to moist soils. During 2020–2021, females tended to have larger home ranges (78.7 ± 46.4 ha) than males (35.0 ± 10.7 ha) and more strongly selected sites closer to riparian corridors, while males selected areas that were closer to upland young forest with flatter slopes than the available surrounding landscape. Such sex-specific differences in habitat selection may be related to males and females prospecting for potential breeding sites during this post-breeding period for the following spring. We used the top-ranked habitat selection models for males and females to produce a spatially explicit state-wide map that identifies low-to-high likelihood of use areas that can be used to guide forest management decisions in southern New England to maximize benefits for American woodcock.     

Effects of crop field characteristics on nocturnal winter use by American woodcock

Since the late 1960s, American woodcock (Scolopax minor) have undergone population declines because of habitat loss. Previous research suggested ridge and furrow topography in conventionally tilled soybean fields provided critical nocturnal cover as birds foraged on earthworms. However, the use of no-till technology has increased and many fields now lack ridge and furrow topography. We assessed woodcock winter nocturnal foraging habitat use given recent changes in agricultural technology, and investigated how field treatment, earthworm abundance, and environmental variables affect the selection of nocturnal foraging sites. We counted woodcock along transects in 5 field treatments twice in each of 67 fields during December–March 2008–2009 and 72 fields during December–March 2009–2010. During both seasons, we collected earthworm and soil samples from a subset of fields of each field treatment. Woodcock densities were at least twice as high in no-till soybean fields planted after corn and in undisked corn fields with mowed stalks than in other field treatments. No-till soybean planted after corn and undisked corn fields contained ridge and furrow topography, whereas other crops did not, and earthworms were at least 1.5 times more abundant in no-till soybean fields than other field treatments. Ridges and furrows in no-till soybean fields planted after corn and undisked corn fields may provide wintering woodcock with thermal protection and concealment from predators. No-till soybean fields planted after corn offered the additional benefit of relatively high food availability. The presence of ridge and furrow topography can be used to predict woodcock field use on the wintering grounds in agricultural areas. Farmers can provide nocturnal winter foraging sites for woodcock by delaying field disking and leaving ridge and furrow topography in crop fields. © 2011 The Wildlife Society.     

American Woodcock Fall Migration Using Central Region Band-Recovery and Wing-Collection Survey Data

ABSTRACT Band-recovery and wing-collection survey data have the potential to provide information on American woodcock (Scolopax minor) fall migration ecology in the Central Region of the United States, yet researchers have not recently analyzed these extensive data sets. We analyzed all direct recoveries of woodcock banded in Michigan, Minnesota, and Wisconsin, USA, as well as wing-collection survey data, to determine the progression of fall migration, the migration direction, and the final destination of woodcock migrating from these states. We did not observe migration initiation based on band recoveries until late October and early November, with most migration occurring during November. Wing-receipt data showed a similar trend, with most change in mean receipt latitude occurring from 1 November to 5 December. During November, wing receipts were spread through the entire Central Region. By 15–31 December, 92% (n = 26) of band recoveries were on the wintering grounds (south of latitude 33°N). Most banded woodcock from Michigan, Minnesota, and Wisconsin wintered in Louisiana, USA. Woodcock banded in these states will be exposed to harvest for most of the hunting season because they remain in these states through November. If the population status of local birds is a concern, managers should consider this migration pattern when setting season dates.     

Migratory connectivity of American woodcock derived using satellite telemetry

American woodcock (Scolopax minor; woodcock) migratory connectivity (i.e., association between breeding and wintering areas) is largely unknown, even though current woodcock management is predicated on such associations. Woodcock are currently managed in the Eastern and Central management regions in the United States with the boundary between management regions analogous to the boundary between the Atlantic and Mississippi flyways, based largely on analysis of band returns from hunters. Factors during migration influence survival and fitness, and existing data derived from banding and very high frequency telemetry provide only coarse-scale information to assess factors influencing woodcock migratory movement patterns and behavior. To assess whether current management-region boundaries correspond with woodcock migratory connectivity in the Central Management Region and to describe migration patterns with higher resolution than has been previously possible, we deployed satellite transmitters on 73 woodcock (25 adult and 28 juvenile females, and 8 adult and 12 juvenile males) and recorded 87 autumn or spring migration paths from 2014 to 2016. Marked woodcock used 2 primary migrations routes: a Western Route and a Central Route. The Western Route ran north-south, connecting the breeding and wintering grounds within the Central Management Region. The hourglass-shaped Central Route connected an area on the wintering grounds reaching from Texas to Florida, to sites throughout northeastern North America in both the Eastern Management Region and Central Management Region and woodcock following this route migrated through the area between the Appalachian Mountains and the Mississippi Alluvial Valley in western Tennessee during autumn and spring. Two of 17 woodcock captured associated with breeding areas in Michigan, Wisconsin, or Minnesota migrated to wintering sites in the Eastern Management Region and 12 marked woodcock captured on wintering areas in Texas and Louisiana migrated to breeding sites in the Eastern Management Region. Woodcock that used the Western Route exhibited high concentrations of stopovers during spring in the Arkansas Ozark Mountains and northern Missouri, and along the Mississippi River on the border between Wisconsin and Minnesota, and autumn concentrations of stopovers in southwestern Iowa, central Missouri, the Arkansas portion of the Ozark Mountains, and around the junction of Texas, Louisiana, Oklahoma, and Arkansas. Woodcock that used the Central Route exhibited high concentrations of stopovers during spring in northern Mississippi through western Tennessee, western Kentucky, and the Missouri Bootheel, and autumn concentrations of stopovers in northern Illinois, southwestern Ohio, and the portions of Kentucky and Tennessee west of the Appalachian Mountains. We suggest that current management of woodcock based on 2 management regions may not be consistent with the apparent lack of strong migratory connectivity we observed. Our results also suggest where management of migration habitat might be most beneficial to woodcock. © 2019 The Wildlife Society.     

Identifying Habitat Linkages for American Black Bears in North Carolina,USA

ABSTRACT Understanding landscape structure and the role of habitat linkages is important to managing wildlife populations in fragmented landscapes. We present a data-based method for identifying local- and regional-scale habitat linkages for American black bears (Ursus americanus) on the Albemarle-Pamlico Peninsula of North Carolina, USA. We used weights-of-evidence, a discrete multivariate technique for combining spatial data, to make predictions about bear habitat use from 1,771 telemetry locations on 2 study areas (n = 35 bears). The model included 3 variables measured at a 0.2-km² scale: forest cohesion, forest diversity, and forest-agriculture edge density, adequately describing important habitat characteristics for bears on our study area. We used 2 categories of unique habitat conditions to delineate favorable bear habitat, which correctly classified 79.5% of the bear locations in a 10-fold model validation. Forest cohesion and forest-agriculture edge density were the most powerful predictors of black bear habitat use. We used predicted probabilities of bear occurrence from the model to delineate habitat linkages among local and regional areas where bear densities were relatively high. Our models clearly identified 2 of the 3 sites previously recommended for wildlife underpasses on a new, 4-lane highway in the study area. Our approach yielded insights into how landscape metrics can be integrated to identify linkages suitable as habitat and dispersal routes.     

The Effect of Local and Landscape-Level Characteristics on the Abundance of Forest Birds in Early-Successional Habitats during the Post-Fledging Season in Western Massachusetts

Many species of mature forest-nesting birds (“forest birds”) undergo a pronounced shift in habitat use during the post-fledging period and move from their forest nesting sites into areas of early-successional vegetation. Mortality is high during this period, thus understanding the resource requirements of post-fledging birds has implications for conservation. Efforts to identify predictors of abundance of forest birds in patches of early-successional habitats have so far been equivocal, yet these previous studies have primarily focused on contiguously forested landscapes and the potential for landscape-scale influences in more fragmented and modified landscapes is largely unknown. Landscape composition can have a strong influence on the abundance and productivity of forest birds during the nesting period, and could therefore affect the number of forest birds in the landscape available to colonize early-successional habitats during the post-fledging period. Therefore, the inclusion of landscape characteristics should increase the explanatory power of models of forest bird abundance in early-successional habitat patches during the post-fledging period. We examined forest bird abundance and body condition in relation to landscape and habitat characteristics of 15 early-successional sites during the post-fledging season in Massachusetts. The abundance of forest birds was influenced by within-patch habitat characteristics, however the explanatory power of these models was significantly increased by the inclusion of landscape fragmentation and the abundance of forest birds in adjacent forest during the nesting period for some species and age groups. Our findings show that including factors beyond the patch scale can explain additional variation in the abundance of forest birds in early-successional habitats during the post-fledging period. We conclude that landscape composition should be considered when siting early-successional habitat to maximize its benefit to forest birds during the post-fledging period, and should also be included in future investigations of post-fledging habitat use by forest birds.     

The effect of habitat on the range expansion of a native and an introduced bird species

Aim Range expansion across a heterogeneous landscape may depend on the habitat selected and used by the expanding species. If habitat selection influences range expansion then localities colonized by a species should contain a greater proportion of favoured habitat (and less non‐habitat) than other nearby localities not colonized. White‐winged doves (Zenaida asiatica) and Eurasian collared doves (Streptopelia decaocto) are two bird species that provide an excellent opportunity to test this hypothesis, because the geographic ranges of both species have been expanding in North America for more than two decades. Location Continental USA. Methods We used distribution data from the North American Breeding Bird Survey to test whether the landscapes occupied by each species contained a greater proportion of favoured habitat (urban land, grassland/pasture, shrub land and cropland) and a lower proportion of non‐habitat (forest land) than landscapes where doves were not found. We tested each species separately in each of three broad expansion areas, namely East, Central and West. We also compared rates of spatial spread between expansion areas and between the two species. Results As predicted, both species tended to occupy landscapes with greater proportions of urban land, shrub land and cropland but with less forest land compared with landscapes without doves, in all three expansion areas. Contrary to prediction, occupied landscapes tended to have slightly less grassland/pasture than unoccupied landscapes. Rates of spread differed between the two species and among expansion areas. Main conclusions Range expansion and the extent to which a species fills or saturates its range are influenced by the habitat ecology of the expanding species. Species colonize localities based on the availability of suitable habitat. However, the role of habitat in a species’ range expansion does depend somewhat on the greater geographical setting. Over large regional and geographical scales, range expansion (rate of spread and saturation) may proceed unevenly, suggesting that range expansion is a very dynamic and context‐specific process.     

Origin and migration of woodcock Scolopax rusticola wintering in Spain

Woodcock Scolopax rusticola is an important game species in Northern Spain, where it is mainly a wintering species. Knowledge about the migration and origin of the woodcock wintering in Spain is relatively sparse, existing to date only qualitative analyses dating more than a decade. From the analyses of ringing recoveries of woodcock wintering in Spain, we evaluate the relative importance of various countries or regions as sources of the woodcock wintering in Spain, an estimate of their migratory route. Our analyses show the Circum-Baltic Region to be the most important breeding area of the woodcock wintering in Spain. Within that area, both Sweden and Western Russia appear to be particularly important. Analyses of the ringing locations of woodcock ringed during migration and recovered in Spain in winter suggest that woodcock wintering in Spain migrate primarily through the South of the Baltic Sea, Germany and France. The proportion of woodcock ringed in different French regions during the postnuptial migration months (October and November) that was subsequently recovered in Spain (mainly through hunting) declined with the proportion of those birds that was recovered in France (also mainly through hunting). We discuss the management implications of these results.     

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.     

Intermediate habitat associations by hybrids may facilitate genetic introgression in a songbird

Hybridization or the interbreeding of genetically discrete populations or species can occur where ranges of genetically distinct units overlap. Golden‐winged warblers Vermivora chrysoptera, a species that has been in steady decline for decades, highlight the potential population‐level consequences of hybridization. A major factor implicated in their decline is hybridization with their sister species, the blue‐winged warbler Vermivora cyanoptera, which has likely been exacerbated by historic and current land‐use practices. We examined habitat associations of golden‐winged and blue‐winged warblers, phenotypic hybrids, and cryptic hybrids (i.e. mismatch between plumage phenotype and genotype as identified by mitochondrial DNA) in an area of relatively recent range overlap and hybridization in northern New York, USA. To explore the robustness of these results, we then compared the patterns from New York with habitat associations from the central Pennsylvanian Appalachian Mountains where blue‐winged warblers either do not occur or are in very low abundance, yet cryptic golden‐winged warbler hybrids are present. From 2008 to 2011, we captured 122 birds in New York and 28 in Pennsylvania and collected blood samples, which we used to determine maternal ancestry. For each bird captured, we measured territory‐level (50‐m radius circles) habitat, and later used remote‐sensing data to quantify habitat on the territories and in surrounding areas (100‐, 250‐, and 500‐m radius circles). In New York, golden‐winged warblers occupied structurally heterogeneous territories surrounded by homogeneously structured, contiguous deciduous forest, far from urban areas. Blue‐winged warblers showed opposite associations, and hybrids’ habitat associations were typically intermediate. In Pennsylvania, the habitat associations of golden‐winged warblers and their cryptic hybrids were remarkably similar to those in New York. These findings suggest that patterns of habitat occupancy by hybrids may promote contact with golden‐winged warblers and thus likely facilitate genetic introgression, even in areas where the parental species are not sympatric.     

Legacy forest structure increases bird diversity and abundance in aging young forests

Many studies have demonstrated the importance of early‐successional forest habitat for breeding bird abundance, composition, and diversity. However, very few studies directly link measures of bird diversity, composition and abundance to measures of forest composition, and structure and their dynamic change over early succession. This study examines the relationships between breeding bird community composition and forest structure in regenerating broadleaf forests of southern New England, USA, separating the influences of ecological succession from retained stand structure. We conducted bird point counts and vegetation surveys across a chronosequence of forest stands that originated between 2 and 24 years previously in shelterwood timber harvests, a silvicultural method of regenerating oak‐mixed broadleaf forests. We distinguish between vegetation variables that relate to condition of forest regeneration and those that reflect legacy stand structure. Using principal components analyses, we confirmed the distinction between regeneration and legacy vegetation variables. We ran regression analysis to test for relationships between bird community variables, including nesting and foraging functional guild abundances, and vegetation variables. We confirmed these relationships with hierarchical partitioning. Our results demonstrate that regenerating and legacy vegetation correlate with bird community variables across stand phases and that the strength with which they drive bird community composition changes with forest succession. While measures of regeneration condition explain bird abundance and diversity variables during late initiation, legacy stand structure explains them during stem exclusion. Canopy cover, ground‐story diversity, and canopy structure diversity are the most powerful and consistent explanatory variables. Our results suggest that leaving varied legacy stand structure to promote habitat heterogeneity in shelterwood harvests contributes to greater bird community diversity. Interestingly, this is particularly important during the structurally depauperate phase of stem exclusion of young regenerating forests.     

Habitat selection and spatiotemporal interactions of a reintroduced mesocarnivore

Habitat quality and quantity are key factors in evaluating the potential for success of a wildlife translocation. However, because of the difficulty or cost of evaluating these factors, habitat assessments may not include valuable information on important habitat attributes including the abundance and distribution of prey, predators, and competitors. Fishers (Pekania pennanti) are one of the most commonly reintroduced carnivores in North America, and they are a species of conservation concern in their western range. We examined the relative importance of landscape features and species interactions in determining habitat use of a reintroduced population of fishers in the southern Cascade Mountains, Washington, USA. We used detections of prey and predators at 134 remote camera stations, remotely sensed forest structure data, and telemetry locations of fishers in a resource selection function to assess the relative importance of prey, predators, and forest structure in fisher habitat selection. Fishers selected habitats based on forest conditions and activity levels of snowshoe hares (Lepus americanus), whereas bobcat (Lynx rufus) and coyote (Canis latrans) activity levels did not directly affect habitat selection. The probability of fisher use increased in older stands, close to recently disturbed stands, and in areas with intermediate levels of hare activity. Bobcat and hare activity levels were positively correlated, and fishers avoided areas with the greatest hare activity, suggesting that fishers may experience a food-safety tradeoff in the study area. Temporal activity patterns in photo detections indicate that fishers may mediate this danger by avoiding bobcats temporally. Our findings suggest that fishers in Washington prefer habitat mosaics of old and recently disturbed stands where they have greater access to resting structures and hares. Management that maintains mosaics of young and old forest across large landscapes is likely to support fisher recovery. Future reintroduction efforts would benefit from an assessment of prey and predator abundance in proposed reintroduction areas before project initiation. © 2019 The Wildlife Society.     

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.     

The consistency and stability of abundance-occupancy relationships in large-scale population dynamics

1. Abundance-occupancy relationships comprise some of the most general and well-explored patterns in macro-ecology. The theory governing these relationships predicts that species will exhibit a positive interspecific and intraspecific relationship between regional occupancy and local abundance. Abundance-occupancy relationships have important implications in using distributional surveys, such as atlases, to understand and document large-scale population dynamics and the consequences of environmental change. A basic need for interpreting such data bases is a better understanding of whether changes in regional occupancy reflect changes in local abundance across species of varying life-history characteristics. 2. Our objective was to test the predictions of the abundance-occupancy rule using two independent data sets, the New York State Breeding Bird Atlas and the North American Breeding Bird Survey. The New York State Breeding Bird Atlas consists of 5332 25-km(2) survey blocks and is one of the first atlases in the USA to be completed for two time periods (1980-85 and 2000-05). The North American Breeding Survey is a large-scale annual survey intended to document the relative abundance and population change of songbirds throughout the USA. 3. We found that regional occupancy was positively correlated with relative abundance across 98 (beta = 0.60 +/- 0.11 SE, P < 0.001, R(2) = 0.60) and 85 species (beta = 0.67 +/- 0.06 SE, P < 0.001, R(2) = 0.57) in two separate time periods. This relationship proved stable over time and was notably consistent between breeding habitat groups and migratory guilds. 4. Between 1980 and 2005, changes in regional occupancy were highly correlated with long-term abundance trend estimates for 75 species (beta = 5.73 +/- 0.24 SE, P < 0.001, R(2) = 0.88). Over a 20-year period, woodland and resident birds showed an increase in occupancy while grassland species showed the greatest decline; these patterns were mirrored by changes in local abundance. 5. Although exceptions existed, we found most changes in occupancy parallel changes in local abundance. These findings support the basic predictions of the abundance-occupancy rule and demonstrate its consistency and stability in species and groups of varying life-history characteristics.     

Bird assemblages of intensively established pine plantations in Coastal Plain Mississippi

Pine (Pinus spp.) plantations are a common type of managed forest in the southeastern United States that may supply habitat for early successional bird species, many of which are declining. To provide information about young pine plantations as bird habitat, we evaluated spring bird presence in 5 combinations of stand establishment practices during years 2 through 5 post-establishment in the Lower Coastal Plain of Mississippi, USA. We detected 38 species with point counts and compared bird metrics among establishment practices using mixed general linear models. Species richness, total relative abundance, and relative abundance of many species were greater by at least a factor of 1.5–4 in the chemical-only establishment practice than mechanically prepared establishment practices, and values declined by about 5–60% within mechanically prepared establishment practices as herbicide intensity increased. Tree and snag retention contributed to avian abundance and richness in the chemical-only establishment practice. Our study, in conjunction with past research, demonstrated the conservation value to early successional bird species of managed pine stands established with tree retention, indicating that silvicultural and wildlife habitat objectives can be met within the range of stand establishment treatments available to managers. © 2012 The Wildlife Society.     

Differential habitat use between demographic states of black bears in managed timber forests

The long-term effects of intensive forest harvest on sensitive demographic stages of the American black bear (Ursus americanus) have been often overlooked. Much of Maine, USA, is covered in forests that are hospitable to bears and commercial timber harvest. To investigate the potential effects of differing intensities of disturbance on black bears, and on females with cubs particularly, we designed a large-scale natural experiment with 197 motion-sensitive camera sites dispersed over representative forest stands in northern and central Maine. Using multi-state occupancy models, we distinguished the overall trends in space use by females with young versus adult bears without young. Forest disturbance at large spatial scales was positively associated with the probability of use for both demographic groups and the availability of hardwood trees was an additional important factor for habitat use by females with young. Our study illustrates the use of motion-sensitive cameras to monitor and understand habitat use by distinct life-history stages of animals living in human-modified landscapes, and results indicate that managers can maintain black bear habitat in areas of active forest harvest by ensuring the availability of hardwood species.