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Patrick D. Keyser

Publications and source records attributed to Patrick D. Keyser.

14 recordsLinked to original sources

Northern bobwhites select for native grasses on working grazing land

Northern bobwhite ( Colinus virginianus ; bobwhite) populations have experienced an 85% decline across most of its range since the 1960s. The most drastic decreases have been in the southeastern United States where biologists attribute the decline to widespread habitat loss, including conversion of native grasslands to tall fescue ( Lolium arundinaceum ) and other exotic cool-season grass (CSG) pasture and hayfields. Including agricultural lands in conservation programs could improve habitat conditions on a regional scale. One working-lands conservation strategy involves the use of native warm-season grasses (NWSGs) rather than cool-season exotic grasses as economically viable cattle forage. To evaluate this management practice for creating bobwhite habitat, we conducted a field study on a bobwhite population in eastern Kentucky, USA, where grazed NWSG and burned NWSG fields were adjacent to exotic grazed and exotic hayed CSG fields. Between April 2019 and October 2022, we used radio telemetry to track bobwhites to evaluate resource selection at second- and third-order scales. Bobwhites used traditionally managed CSGs less than would be expected by chance at the second-order scale in both breeding and non-breeding seasons. Bobwhite use of grazed NWSG and burned NWSG was greater than would be expected by chance at the second-order scale in both breeding and non-breeding seasons. Similarly, at the third-order scale, bobwhites used CSGs less than would be expected by chance, whereas NWSGs and woody stems were used more than by chance alone. Our study suggests that bobwhites select NWSGs at multiple spatial scales despite broad-scale availability of managed exotic CSGs. Under a working-lands framework, the integration of NWSGs into working lands could create bobwhite habitat in the southeastern United States, especially if woody stems are present.

Kentucky↗

Northern bobwhite breeding season ecology on a reclaimed surface mine

Surface coal mining and subsequent reclamation of surface mines have converted large forest areas into early successional vegetative communities in the eastern United States. This reclamation can provide a novel opportunity to conserve northern bobwhite ( Colinus virginianus ). We evaluated the influence of habitat management activities on nest survival, nest-site selection, and brood resource selection on managed and unmanaged units of a reclaimed surface mine, Peabody Wildlife Management Area (Peabody), in west-central Kentucky, USA, from 2010 to 2013. We compared resource selection, using discrete-choice analysis, and nest survival, using the nest survival model in Program MARK, between managed and unmanaged units of Peabody at 2 spatial scales: the composition and configuration of vegetation types (i.e., macrohabitat) and vegetation characteristics at nest sites and brood locations (i.e., microhabitat). On managed sites, we also investigated resource selection relative to a number of different treatments (e.g., herbicide, disking, prescribed fire). We found no evidence that nest-site selection was influenced by macrohabitat variables, but bobwhite selected nest sites in areas with greater litter depth than was available at random sites. On managed units, bobwhite were more likely to nest where herbicide was applied to reduce sericea lespedeza ( Lespedeza cuneata ) compared with areas untreated with herbicide. Daily nest survival was not influenced by habitat characteristics or by habitat management but was influenced by nest age and the interaction of nest initiation date and nest age. Daily nest survival was greater for older nests occurring early in the breeding season (0.99, SE < 0.01) but was lower for older nests occurring later in the season (0.08, SE = 0.13). Brood resource selection was not influenced by macrohabitat or microhabitat variables we measured, but broods on managed units selected areas treated with herbicide to control sericea lespedeza and were located closer to firebreaks and disked native-warm season grass stands than would be expected at random. Our results suggest the vegetation at Peabody was sufficient without manipulation to support nesting and brood-rearing northern bobwhite at a low level, but habitat management practices improved vegetation for nesting and brood-rearing resource selection. Reproductive rates (e.g., nest survival and re-nesting rates) at Peabody were lower than reported in other studies, which may be related to nutritional deficiencies caused by the abundance of sericea lespedeza. On reclaimed mine lands dominated by sericea lespedeza, we suggest continuing practices such as disking and herbicide application that are targeted at reducing sericea lespedeza to improve the vegetation for nesting and brood-rearing bobwhite.

Kentucky↗

Habitat manipulation influences northern bobwhite resource selection on a reclaimed surface mine

More than 600,000&thinsp;ha of mine land have been reclaimed in the eastern United States, providing large contiguous tracts of early successional vegetation that can be managed for northern bobwhite ( Colinus virginianus ). However, habitat quality on reclaimed mine land can be limited by extensive coverage of non-native invasive species, which are commonly planted during reclamation. We used discrete-choice analysis to investigate bobwhite resource selection throughout the year on Peabody Wildlife Management Area, a 3,330-ha reclaimed surface mine in western Kentucky. We used a treatment-control design to study resource selection at 2 spatial scales to identify important aspects of mine land vegetation and whether resource selection differed between areas with habitat management (i.e., burning, disking, herbicide; treatment) and unmanaged units (control). Our objectives were to estimate bobwhite resource selection on reclaimed mine land and to estimate the influence of habitat management practices on resource selection. We used locations from 283 individuals during the breeding season (1 Apr&ndash;30 Sep) and 136 coveys during the non-breeding season (1 Oct&ndash;Mar 31) from August 2009 to March 2014. Individuals were located closer to shrub cover than would be expected at random throughout the year. During the breeding season, individuals on treatment units used areas with smaller contagion index values (i.e., greater interspersion) compared with individuals on control units. During the non-breeding season, birds selected areas with greater shrub-open edge density compared with random. At the microhabitat scale, individuals selected areas with increased visual obstruction >1&thinsp;m aboveground. During the breeding season, birds were closer to disked areas (linear and non-linear) than would be expected at random. Individuals selected non-linear disked areas during winter but did not select linear disked areas (firebreaks) because they were planted to winter wheat each fall and lacked cover during the non-breeding season. Individuals also selected areas treated with herbicide to control sericea lespedeza ( Lespedeza cuneata ) throughout the year. During the breeding season, bobwhites avoided areas burned during the previous dormant season. Habitat quality of reclaimed mine lands may be limited by a lack of shrub cover and extensive coverage of non-native herbaceous vegetation. Managers aiming to increase bobwhite abundance should focus on increasing interspersion of shrub cover, with no area >100&thinsp;m from shrub cover. We suggest disking and herbicide application to control invasive species and improve the structure and composition of vegetation for bobwhites.

Kentucky↗

Impact of experimental habitat manipulation on northern bobwhite survival

Habitat management for northern bobwhite ( Colinus virginianus ) should affect vital rates, but direct linkages with survival are not well documented; therefore, we implemented an experiment to evaluate those responses. We conducted our experiment on a reclaimed surface mine, a novel landscape where conditions were considered sub-optimal because of the dominance of non-native vegetation, such as sericea lespedeza ( Lespedeza cuneata ), which has been reported to provide marginal habitat for northern bobwhite and may negatively affect survival. Nonetheless, these areas have great potential for contributing to bobwhite conservation because of the amount of early successional cover they provide. Our study site, a 3,330-ha reclaimed surface mine in western Kentucky, consisted of 2 tracts (Sinclair and Ken, 1,471&thinsp;ha and 1,853&thinsp;ha, respectively) that served as replicates with each randomly divided into a treatment (i.e., habitat manipulation through a combination of disking, burning, and herbicide application) and an undisturbed control ( n &thinsp;=&thinsp;4 experimental units). Habitat treatments were applied October 2009 to September 2013. We used radio telemetry to monitor northern bobwhite ( n &thinsp;=&thinsp;1,198) during summer (1 Apr&ndash;30 Sep) and winter (1 Oct&ndash;31 Mar), 2009&ndash;2013. We used the known-fate model in Program MARK to evaluate treatment effects on seasonal survival rates. We included biological, home-range, landscape, and microhabitat metrics as covariates to help improve model sensitivity and further elucidate experimental impacts. Survival varied annually, ranging from 0.139 (SE&thinsp;=&thinsp;0.031) to 0.301 (SE&thinsp;=&thinsp;0.032), and seasonally (summer, 0.148 [SE&thinsp;=&thinsp;0.015]; winter, 0.281 [SE&thinsp;=&thinsp;0.022]). We found a treatment effect ( &beta; &thinsp;=&thinsp;0.256, 95% CI&thinsp;=&thinsp;0.057&ndash;0.456) with a seasonal interaction ( &beta; &thinsp; =&thinsp;&minus;0.598, 95% CI&thinsp;=&thinsp;&minus;0.898 to &minus;0.298) with survival being higher in summer (0.179 [SE&thinsp;=&thinsp;0.022] vs. 0.109 [SE&thinsp;=&thinsp;0.019]) and lower in winter (0.233 [SE&thinsp;=&thinsp;0.025] vs. 0.355 [SE&thinsp;=&thinsp;0.035]) on treatment than control units. Among habitat covariates, litter depth ( &beta; &thinsp;=&thinsp;&minus;0.387, 95%CI&thinsp;=&thinsp;&minus;0.5809 to &minus;0.1930) was the most influential effect (negative) on survival. Additional experiments across a wider range of habitat conditions may be required to determine management intensity or duration thresholds required to elicit greater changes in survival for northern bobwhite populations.

Kentucky↗

Habitat quality from individual- and population-level perspectives and implications for management

Many wildlife management prescriptions are either implicitly or explicitly designed to improve habitat quality for a focal species, but habitat quality is often difficult to quantify. Depending upon the approach used to define and identify high-quality habitat, management decisions may differ widely. Although individual-level measures of habitat quality based on per capita reproduction (e.g., average nesting success, number of young produced per pair) are most common in the literature, they may not align with population-level measures that reflect number of young produced within a defined area. Using data on the cerulean warbler ( Setophaga cerulea ) collected in the Cumberland Mountains (Tennessee, USA; 2008&ndash;2010) as an example, we illustrate how lack of concordance between individual- and population-level measures of habitat quality can have real-world management implications.

Wildlife Society Bulletin↗

Effects of population reduction on white-tailed deer home-range dynamics

Management strategies designed to reduce the negative impacts of overabundant Odocoileus virginianus (White-tailed Deer) populations on forest regeneration may be influenced by changes in both population density and timber harvest. However, there is conflicting evidence as to how such changes in per capita resource availability influence home-range patterns. We compared home-range patterns of 33 female White-tailed Deer from a low-density population at a site with abundant browse to patterns of a sample of >100 females prior to a 75% reduction in population density and a doubling in timber harvest area. Home-range and core-area sizes were approximately 3 times larger than were found prior to population decline and timber harvest increase, consistent with predictions related to intraspecific competition. We also observed greater site fidelity than previously exhibited, although this may be an artifact of increased home-range sizes. Our results support previous research suggesting that White-tailed Deer home-range size is inversely related to population density and is driven, in part, by intraspecific competition for resources. Relationships among population density, resource availability, and home-range patterns among female White-tailed Deer appear to be complex and context specific.

West Virginia↗

Avian response to timber harvesting applied experimentally to manage Cerulean Warbler breeding populations

Timber harvesting has been proposed as a management tool to enhance breeding habitat for the Cerulean Warbler ( Setophaga cerulea ), a declining Neotropical&ndash;Nearctic migratory songbird that nests in the canopy of mature eastern deciduous forests. To evaluate how this single-species management focus might fit within an ecologically based management approach for multiple forest birds, we performed a manipulative experiment using four treatments (three intensities of timber harvests and an unharvested control) at each of seven study areas within the core Cerulean Warbler breeding range. We collected pre-harvest (one year) and post-harvest (four years) data on the territory density of Cerulean Warblers and six additional focal species, avian community relative abundance, and several key habitat variables. We evaluated the avian and habitat responses across the 3&ndash;32 m 2 ha &minus;1 residual basal area (RBA) range of the treatments. Cerulean Warbler territory density peaked with medium RBA (&sim;16 m 2 ha &minus;1 ). In contrast, territory densities of the other focal species were negatively related to RBA (e.g., Hooded Warbler [ Setophaga citrina ]), were positively related to RBA (e.g., Ovenbird [ Seiurus aurocapilla ]), or were not sensitive to this measure (Scarlet Tanager [ Piranga olivacea ]). Some species (e.g., Hooded Warbler) increased with time post-treatment and were likely tied to a developing understory, whereas declines (e.g., Ovenbird) were immediate. Relative abundance responses of additional species were consistent with the territory density responses of the focal species. Across the RBA gradient, greatest separation in the avian community was between early successional forest species (e.g., Yellow-breasted Chat [ Icteria virens ]) and closed-canopy mature forest species (e.g., Ovenbird), with the Cerulean Warbler and other species located intermediate to these two extremes. Overall, our results suggest that harvests within 10&ndash;20 m 2 ha &minus;1 RBA yield the largest increases in Cerulean Warblers, benefit additional disturbance-dependent species, and may retain closed-canopy species but at reduced levels. Harvests outside the optimum RBA range for Cerulean Warblers can support bird assemblages specifically associated with early or late (closed-canopy) successional stages.

Forest Ecology and Management↗

Multiple plumage traits convey information about age and within-age-class qualities of a canopy-dwelling songbird, the Cerulean Warbler

Colorful plumage traits in birds may convey multiple, redundant, or unreliable messages about an individual. Plumage may reliably convey information about disparate qualities such as age, condition, and parental ability because discrete tracts of feathers may cause individuals to incur different intrinsic or extrinsic costs. Few studies have examined the information content of plumage in a species that inhabits forest canopies, a habitat with unique light environments and selective pressures. We investigated the information content of four plumage patches (blue-green crown and rump, tail white, and black breast band) in a canopy-dwelling species, the Cerulean Warbler ( Setophaga cerulea ), in relation to age, condition, provisioning, and reproduction. We found that older males displayed wider breast bands, greater tail white, and crown and rump feathers with greater blue-green (435&ndash;534 nm) chroma and hue than males in their first potential breeding season. In turn, older birds were in better condition (short and long term) and were reproductively superior to younger birds. We propose that these age-related plumage differences (i.e. delayed plumage maturation) were not a consequence of a life history strategy but instead resulted from constraints during early feather molts. Within age classes, we found evidence to support the multiple messages hypothesis. Birds with greater tail white molted tails in faster, those with more exaggerated rump plumage (lower hue, greater blue-green chroma) provisioned more, and those with lower rump blue-green chroma were in better condition. Despite evidence of reliable signaling in this species, we found no strong relationships between plumage and reproductive performance, potentially because factors other than individual differences more strongly influenced fecundity.

The Auk↗

Comparison of point counts and territory mapping for detecting effects of forest management on songbirds

Point counts are commonly used to assess changes in bird abundance, including analytical approaches such as distance sampling that estimate density. Point-count methods have come under increasing scrutiny because effects of detection probability and field error are difficult to quantify. For seven forest songbirds, we compared fixed-radii counts (50 m and 100 m) and density estimates obtained from distance sampling to known numbers of birds determined by territory mapping. We applied point-count analytic approaches to a typical forest management question and compared results to those obtained by territory mapping. We used a before&ndash;after control impact (BACI) analysis with a data set collected across seven study areas in the central Appalachians from 2006 to 2010. Using a 50-m fixed radius, variance in error was at least 1.5 times that of the other methods, whereas a 100-m fixed radius underestimated actual density by >3 territories per 10 ha for the most abundant species. Distance sampling improved accuracy and precision compared to fixed-radius counts, although estimates were affected by birds counted outside 10-ha units. In the BACI analysis, territory mapping detected an overall treatment effect for five of the seven species, and effects were generally consistent each year. In contrast, all point-count methods failed to detect two treatment effects due to variance and error in annual estimates. Overall, our results highlight the need for adequate sample sizes to reduce variance, and skilled observers to reduce the level of error in point-count data. Ultimately, the advantages and disadvantages of different survey methods should be considered in the context of overall study design and objectives, allowing for trade-offs among effort, accuracy, and power to detect treatment effects.

Journal of Field Ornithology↗

Emulating natural disturbances for declining late-successional species: A case study of the consequences for Cerulean Warblers ( Setophaga cerulea )

Forest cover in the eastern United States has increased over the past century and while some late-successional species have benefited from this process as expected, others have experienced population declines. These declines may be in part related to contemporary reductions in small-scale forest interior disturbances such as fire, windthrow, and treefalls. To mitigate the negative impacts of disturbance alteration and suppression on some late-successional species, strategies that emulate natural disturbance regimes are often advocated, but large-scale evaluations of these practices are rare. Here, we assessed the consequences of experimental disturbance (using partial timber harvest) on a severely declining late-successional species, the cerulean warbler ( Setophaga cerulea ), across the core of its breeding range in the Appalachian Mountains. We measured numerical (density), physiological (body condition), and demographic (age structure and reproduction) responses to three levels of disturbance and explored the potential impacts of disturbance on source-sink dynamics. Breeding densities of warblers increased one to four years after all canopy disturbances (vs. controls) and males occupying territories on treatment plots were in better condition than those on control plots. However, these beneficial effects of disturbance did not correspond to improvements in reproduction; nest success was lower on all treatment plots than on control plots in the southern region and marginally lower on light disturbance plots in the northern region. Our data suggest that only habitats in the southern region acted as sources, and interior disturbances in this region have the potential to create ecological traps at a local scale, but sources when viewed at broader scales. Thus, cerulean warblers would likely benefit from management that strikes a landscape-level balance between emulating natural disturbances in order to attract individuals into areas where current structure is inappropriate, and limiting anthropogenic disturbance in forests that already possess appropriate structural attributes in order to maintain maximum productivity.

PLoS ONE↗

Spatial variation in breeding habitat selection by Cerulean Warblers ( Setophaga cerulea ) throughout the Appalachian Mountains

Studies of habitat selection are often of limited utility because they focus on small geographic areas, fail to examine behavior at multiple scales, or lack an assessment of the fitness consequences of habitat decisions. These limitations can hamper the identification of successful site-specific management strategies, which are urgently needed for severely declining species like Cerulean Warblers ( Setophaga cerulea ). We assessed how breeding habitat decisions made by Cerulean Warblers at multiple scales, and the subsequent effects of these decisions on nest survival, varied across the Appalachian Mountains. Selection for structural habitat features varied substantially among areas, particularly at the territory scale. Males within the least-forested landscapes selected microhabitat features that reflected more closed-canopy forest conditions, whereas males in highly forested landscapes favored features associated with canopy disturbance. Selection of nest-patch and nest-site attributes by females was more consistent across areas, with females selecting for increased tree size and understory cover and decreased basal area and midstory cover. Floristic preferences were similar across study areas: White Oak ( Quercus alba ), Cucumber-tree ( Magnolia acuminata ), and Sugar Maple ( Acer saccharum ) were preferred as nest trees, whereas red oak species (subgenus Erythrobalanus ) and Red Maple ( A. rubrum ) were avoided. The habitat features that were related to nest survival also varied among study areas, and preferred features were negatively associated with nest survival at one area. Thus, our results indicate that large-scale spatial heterogeneity may influence local habitat-selection behavior and that it may be necessary to articulate site-specific management strategies for Cerulean Warblers.

Ohio, Kentucky, Tennessee, West Virginia↗

Management guidelines for enhancing Cerulean Warbler breeding habitat in Appalachian hardwood forests

The Cerulean Warbler (Setophaga cerulea) is a migratory songbird that breeds in mature deciduous forests of eastern North America. Cerulean Warblers (hereafter, ceruleans) require heavily forested landscapes for nesting and, within Appalachian forests, primarily occur on ridge tops and steep, upper slopes. They are generally associated with oakdominated (Quercus spp.) stands that contain gaps in the forest canopy, that have large diameter trees (>16 inches diameter breast height (dbh)), and that have well-developed understory-and upper-canopy layers. Ceruleans primarily use the midand upper-canopy where they glean insects from the surface of leaves and conceal their open cup nests. Because they are severely declining across much of their range (Fig. 1), habitat management is a high priority. Management for this species can also improve conditions for a number of other wildlife species that depend on the same structure.

Appalachian Mountains↗

Canada Warbler use of harvested stands following timber management in the southern portion of their range

We evaluated Canada Warbler ( Cardellina canadensis ) responses to changes in habitat characteristics (landscape metrics, landcover, and microhabitat features) at multiple spatial scales resulting from timber harvests (clear-cut, heavy partial, and light partial) between 1996 and 2009. Relative abundance of Canada Warblers decreased over time on our West Virginia study area (Wildlife and Ecosystem Research Forest) and within the Appalachian Bird Conservation Region. Initially, relative abundance was greater closer to roads, but as timber harvests became more common, relative abundance became positively associated with amount of light partial harvests at the local scale. Nest survival was 45.6 ± 18.3% during 1996–1998 and 24.9 ± 14.6% during 2007–2009, but did not differ ( P = 0.38) between these periods. Areas around nests in 2007–2009 ( n = 17) had less intermediate canopy cover and fewer residual trees but more green cover, woody debris, and pole trees than areas around nests in 1996–1998 ( n = 10). Successful nests had more low cover, less vertical diversity, more woody debris, more saplings, and greater edge density than failed nests. We found a positive association between relative abundance and all three types of timber harvests and an improvement in habitat through understory development and retention of residual trees. Our research finds preliminary support for use of timber harvests, particularly light partial harvests, as a management tool for Canada Warblers in the southern portion of their range with the need for extended research using treatments and controls to confirm successful management.

Forest Ecology and Management↗

Threshold responses of songbirds to long-term timber management on an active industrial forest

Forest managers often seek to balance economic benefits from timber harvesting with maintenance of habitat for wildlife, ecosystem function, and human uses. Most research on the relationship between avian abundance and active timber management has been short-term, lasting one to two years, creating the need to investigate long-term avian responses and to identify harvest thresholds when a small change in habitat results in a disproportionate response in relative abundance and nest success. Our objectives were to identify trends in relative abundance and nest success and to identify landscape-scale disturbance thresholds for avian species and habitat guilds in response to a variety of harvest treatments (clear-cuts, heavy and light partial harvests) over 14 years. We conducted point counts and monitored nests at an industrial forest in the central Appalachians of West Virginia during 1996–1998, 2001–2003, and 2007–2009. Early successional species increased in relative abundance across all three time periods, whereas interior-edge and forest-interior guilds peaked in relative abundance mid-study after which the forest-interior guild declined. Of 41 species with >10 detections, four (10%) declined significantly, 13 (32%) increased significantly (only three species among all periods), and 9 (22%) peaked in abundance mid-study (over the entire study period, four species had no significant change in abundance, four declined, and one increased). Based on piecewise linear models, forest-interior and interior-edge guilds’ relative abundance harvest thresholds were 28% total harvests (all harvests combined), 10% clear-cut harvests, and 18% light partial harvests, after which abundances declined. Harvest thresholds for the early successional guild were 42% total harvests, 11% clear-cut harvest, and 10% light partial harvests, and relative abundances increased after surpassing thresholds albeit at a reduced rate of increase after the clear-cut threshold. Threshold confidence intervals for individual species overlapped their guild threshold intervals 91% of the time. Even though relative abundance of most species (80%) did not decline as the area affected by timber management increased, implementing management at or below our approximate forest-interior and interior-edge harvest thresholds would reduce the number of declining species by half, maintain higher relative abundances of four species with a net decline in abundance but that peaked in abundance mid-study, and maintain higher relative abundances of ten additional species. In contrast, this management strategy also would prevent the increase in relative abundance of seven species and limit the increase in abundance of three species that increased throughout the study.

West Virginia↗