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Experimental analysis and simulation modeling of forest management impacts on wood thrushes, Hylocichla mustelina

North American Breeding Bird Survey data show that wood thrush (Hylocichla mustelina) populations in eastern U.S. forests have declined 1.8% per year during 1966-95. The declining quality of breeding forest tracts in North America is one possible cause for the apparent decline of some neotropical migratory birds, such as the wood thrush. In Georgia, however, wood thrush populations have declined during a period of increasing pine forest area and larger patch sizes. We hypothesized that forest management practices such as thinning and prescribed burning might create unsuitable habitat for wood thrushes. We conducted a four-year before/after, treatment/control experiment at the Piedmont National Wildlife Refuge in central Georgia to study to the effects of a treatment of thinning and prescribed burning on wood thrush demographic parameters. We simultaneously monitored wood thrush adults and juveniles with mark-recapture, radio-telemetry, nest searches, and plot-map surveys. Our analyses showed that wood thrushes were less likely to emigrate from the study compartments after the treatment, and wood thrushes exhibited some tendency to increase preference for hardwood habitats and decrease preference for pine habitats following the treatment. However, we observed no effects of treatment on nest success, adult survival, and adult and juvenile dispersal distances. We also found that female wood thrushes had lower survival rates than males during the breeding season, and we documented large-scale, within-year dispersal movements of adult (up to 17 km) and juvenile (up to 7 km) wood thrushes. We conclude that landscape level habitat quantity and quality must be considered during songbird management decisions. The documentation of sex- and age-specific wood thrush survival and movement rates was critical for construction of a set of population models. We used three stochastic models to learn more about wood thrush population dynamics and make predictions about population growth rates, reproductive success, and the effect of habitat changes on wood thrush populations. The simplest source/sink population model suggests that the Piedmont National Wildlife Refuge's wood thrush population is probably stable or increasing, and wood thrush populations in treated areas had higher growth rates than birds in untreated areas. We were able to use the individual-based model of wood thrush productivity to predict fecundity, a parameter that we could not measure directly in the field. Again, females on treated areas had higher fecundity than birds on untreated areas. Our spatially-based model predicted that wood thrush populations should respond positively to predicted changes in the age/size class structure of the Refuge's pine forests. Our model also showed that most wood thrushes leave the Refuge's forest compartments during the breeding season, and these dispersal movements are extremely important to understanding and managing wood thrush populations. The use of prescribed burning and retention shelterwood silviculture at the Piedmont National Wildlife Refuge does not appear to negatively affect the local wood thrush population. Continued use of the current management regime should result in adequate nesting, foraging, and escape habitats for wood thrushes. However, landscape-level habitat availability and quality, including lands outside the Refuge, must be considered when making management decisions that may affect wood thrushes.

Book

Effects of radio transmitters on migrating wood thrushes

We quantified the effects of radio transmitters on Wood Thrushes (Hylocichla mustelina) using 4 yr of banding and telemetry data from Piedmont National Wildlife Refuge, Georgia. Flight performance models suggest that the 1.6-g transmitter shortens the migratory range of Wood Thrushes by only 60 km, and the estimated migratory range is adequate to accomplish migration even with limited fat stores. We used two strengths of line, 5- and 9-kg test-strength braided Dacron, to attach the transmitters using the thigh-harness method. We recaptured 13 returning radio-marked Wood Thrushes, seven of which were still marked. Six of the seven birds marked with the 5-kg test harnesses lost their transmitters within 1 yr while all six of the 9-kg test harnesses were still attached up to 21 mo later. Radio-marking did not reduce the return rates of adults and immatures, and the transmitters did not cause radio-marked birds to lose more mass than banded-only birds. Wood Thrushes can successfully carry a transmitter during migration with no detectable negative effects. We recommend continued use of the thigh-harness method, but we encourage the use of 5-kg cotton line.

Journal of Field Ornithology

Determinants of wood thrush nest success: A multi-scale, model selection approach

We collected data on 212 wood thrush (Hylocichla mustelina) nests in central New York from 1998 to 2000 to determine the factors that most strongly influence nest success. We used an information-theoretic approach to assess and rank 9 models that examined the relationship between nest success (i.e., the probability that a nest would successfully fledge at least 1 wood thrush offspring) and habitat conditions at different spatial scales. We found that 4 variables were significant predictors of nesting success for wood thrushes: (1) total core habitat within 5 km of a study site, (2) distance to forest-field edge, (3) total forest cover within 5 km of the study site, and (4) density and variation in diameter of trees and shrubs surrounding the nest. The coefficients of these predictors were all positive. Of the 9 models evaluated, amount of core habitat in the 5-km landscape was the best-fit model, but the vegetation structure model (i.e., the density of trees and stems surrounding a nest) was also supported by the data. Based on AIC weights, enhancement of core area is likely to be a more effective management option than any other habitat-management options explored in this study. Bootstrap analysis generally confirmed these results; core and vegetation structure models were ranked 1, 2, or 3 in over 50% of 1,000 bootstrap trials. However, bootstrap results did not point to a decisive model, which suggests that multiple habitat factors are influencing wood thrush nesting success. Due to model uncertainty, we used a model averaging approach to predict the success or failure of each nest in our dataset. This averaged model was able to correctly predict 61.1% of nest outcomes.

Journal of Wildlife Management

Wood Thrush movements and habitat use: Effects of forest management for Red-cockaded Woodpeckers

We monitored adult and juvenile breeding-season movements and habitat use of radio-tagged Wood Thrushes ( Hylocichla mustelina ) at the Piedmont National Wildlife Refuge, central Georgia, USA. We investigated the effects that management for Red-cockaded Woodpeckers ( Picoides borealis ), thinning and burning >30 year old loblolly pine ( Pinus taeda ) habitat, had on Wood Thrushes, a ground-foraging and midstory-nesting species. Adult Wood Thrush pairs regularly moved long distances between nesting attempts (range 1 to 17,388 m). The only experimental effect we found on adult movements was a decrease in weekly emigration rates (Ψ) from thinned and burned compartments after silvicultural management. Adult males preferred riparian hardwoods with sparse to moderate cover and those preferences increased following management. Juveniles remained near their nest site ( x̄ = 177 m, SE = 113) for an average 24 days (SE = 6.3), and then dispersed a mean 2,189 m (SE = 342). Before dispersal, juveniles preferred upland hardwood–pine mixed habitat ( P < 0.05) with moderate overstory cover ( P < 0.05). We found no management effects on dispersal distances or predispersal habitat use. However, juveniles from thinned and burned compartments dispersed to hardwood habitats with dense cover, whereas birds from control compartments dispersed to pine-dominated habitats with sparse cover. All juveniles dispersed to areas with habitat similar to what they used before dispersal. Small-scale thinning and burning for Red-cockaded Woodpeckers may have had little effect on Wood Thrush habitat use and movements because typical movements were often larger than the scale (stand or compartment) targeted for management.

The Auk

Effects of forest management on density, survival, and population growth of wood thrushes

Loss and alteration of breeding habitat have been proposed as causes of declines in several Neotropical migrant bird populations. We conducted a 4-year study to determine the effects of winter prescribed burning and forest thinning on breeding wood thrush ( Hylocichla mustelina ) populations at the Piedmont National Wildlife Refuge (PNWR) in Georgia. We estimated density, adult and juvenile survival rates, and apparent annual survival using transect surveys, radiotelemetry, and mist netting. Burning and thinning did not cause lower densities ( P = 0.25); wood thrush density ranged from 0.15 to 1.30 pairs/10 ha. No radiomarked male wood thrushes ( n = 68) died during the 4 years, but female ( n = 63) weekly survival was 0.981 ± 0.014 (SE) for females ( n = 63) and 0.976 ± 0.010 for juveniles ( n = 38). Apparent annual adult survival was 0.579 (SE = 0.173). Thinning and prescribed burning did not reduce adult or juvenile survival during the breeding season or apparent annual adult survival. Annual population growth (lambda) at PNWR was 1.00 (95% confidence interval = 0.32-1.63), and the considerable uncertainty in this prediction underscores the need for long term monitoring to effectively manage Neotropical migrants. Population growth increased on experimental compartments after the burn and thin (95% CI before = 0.91-0.97, after = 0.98-1.05), while control compartment declined (before = 0.98-1.05, after = 0.87-0.92). We found no evidence that the current management regime at PNWR, designed to improve red-cockaded woodpecker ( Picoides borealis ) habitat, negatively affected wood thrushes.

Georgia

Effects of early-successional shrubland management on breeding wood thrush populations

In forested landscapes, creation of habitat for early-successional shrubland birds is controversial because of perceived conflicts with the conservation of mature-forest birds. Nonetheless, many mature-forest birds, especially fledglings, readily use early-successional stands during the post-breeding period. This suggests that for mature-forest birds, creating habitat for early-successional birds could involve a tradeoff: reduced abundance and nest survival due to the loss of nesting habitat versus enhanced fledgling survival in early-successional stands. Our research addressed the effects of the creation of early-successional habitat for shrubland birds on wood thrushes ( Hylocichla mustelina ) in western Massachusetts, USA. We compared wood thrush abundance, nest success, fecundity, and post-fledging survival in landscapes with high (∼20%) or low (∼1%) cover of early-successional stands suitable for shrubland birds. We found no differences in nest success, fecundity, and post-fledging survival between the 2 types of landscapes. Abundance of breeders, however, was significantly greater on the sites with high cover of early-successional habitat. We conclude that in forested landscapes, creation of early-successional habitat at levels recommended for the conservation of shrubland birds is compatible with viable wood thrush populations. © 2018 The Wildlife Society.

Massachusetts

Are traditional methods of determining nest predators and nest fates reliable? An experiment with Wood Thrushes ( Hylocichla mustelina ) using miniature video cameras

We used miniature infrared video cameras to monitor Wood Thrush ( Hylocichla mustelina ) nests during 1998–2000. We documented nest predators and examined whether evidence at nests can be used to predict predator identities and nest fates. Fifty-six nests were monitored; 26 failed, with 3 abandoned and 23 depredated. We predicted predator class (avian, mammalian, snake) prior to review of video footage and were incorrect 57% of the time. Birds and mammals were underrepresented whereas snakes were over-represented in our predictions. We documented ≥9 nest-predator species, with the southern flying squirrel ( Glaucomys volans ) taking the most nests ( n = 8). During 2000, we predicted fate (fledge or fail) of 27 nests; 23 were classified correctly. Traditional methods of monitoring nests appear to be effective for classifying success or failure of nests, but ineffective at classifying nest predators.

The Auk

Landscape context moderates edge effects: Nesting success of wood thrushes in Central New York

Despite two decades of research into the effects of habitat fragmentation and edges on nesting birds, critical information about how edges affect the success of natural nests of Neotropical migratory songbirds breeding in heterogeneous landscapes is still missing. We studied abundance and nesting success in Wood Thrushes (Hylocichla mustelina) breeding across a heterogeneous landscape in central New York from 1998 to 2000 to test the hypothesis that edge effects on nesting passerines are stronger in fragmented than contiguous landscapes. We monitored nests to estimate nesting success in edge and interior habitats in both fragmented and contiguously forested landscapes. In contiguous landscapes, daily survival rate did not differ between edge nests (0.963) and interior nests (0.968) (??2 = 0.19, p = 0.66). In contrast, in fragmented landscapes, daily survival estimates were higher in interior (0.971) than edge (0.953) nests (??2 = 3.1, p = 0.08). Our study supports the hypothesis that landscape composition moderates edge effects on actual nests of birds but does not determine the mechanisms causing these patterns.

Conservation Biology

Fat, weather, and date affect migratory songbirds’ departure decisions, routes, and time it takes to cross the Gulf of Mexico

Approximately two thirds of migratory songbirds in eastern North America negotiate the Gulf of Mexico (GOM), where inclement weather coupled with no refueling or resting opportunities can be lethal. However, decisions made when navigating such features and their consequences remain largely unknown due to technological limitations of tracking small animals over large areas. We used automated radio telemetry to track three songbird species (Red-eyed Vireo, Swainson&rsquo;s Thrush, Wood Thrush) from coastal Alabama to the northern Yucatan Peninsula (YP) during fall migration. Detecting songbirds after crossing &sim;1,000 km of open water allowed us to examine intrinsic (age, wing length, fat) and extrinsic (weather, date) variables shaping departure decisions, arrival at the YP, and crossing times. Large fat reserves and low humidity, indicative of beneficial synoptic weather patterns, favored southward departure across the Gulf. Individuals detected in the YP departed with large fat reserves and later in the fall with profitable winds, and flight durations (mean = 22.4 h) were positively related to wind profit. Age was not related to departure behavior, arrival, or travel time. However, vireos negotiated the GOM differently than thrushes, including different departure decisions, lower probability of detection in the YP, and longer crossing times. Defense of winter territories by thrushes but not vireos and species-specific foraging habits may explain the divergent migratory behaviors. Fat reserves appear extremely important to departure decisions and arrival in the YP. As habitat along the GOM is degraded, birds may be limited in their ability to acquire fat to cross the Gulf.

Proceedings of the National Academy of Sciences

Occurrence and nest survival of four thrush species on a managed central Appalachian forest

The wood thrush (Hylocichla mustelina Gmelin) is a species of concern in the central Appalachians, and is sympatric there with three related species, the American robin (Turdus migratorius Linnaeus), hermit thrush (Catharus guttatus Pallas), and veery (Catharus fuscescens Stephens). Our objectives were to quantify use of mature forests and areas subjected to even-aged harvesting and partial harvesting by these four species by measuring their frequency of occurrence, nest survival, and nest site characteristics. We also compared microhabitat characteristics among the landcover types. During 2001-2003 we conducted point count surveys, monitored nests, and collected nest habitat data on a managed forest in West Virginia. Land cover was digitized into five categories: deciduous and mixed mature forest, deciduous and mixed partial harvest, and even-aged regeneration harvest. Chi-square goodness-of-fit analysis with Bonferroni 95% confidence intervals indicated that deciduous partial harvests were more likely to be inhabited by wood thrushes. The other three species were less likely to occur in deciduous partial harvests, and veery had lower nest survival in partial harvests than in mature forest. Contrary to many published descriptions that suggest thrushes will not nest in even-aged harvests, a small number of all species but hermit thrushes did nest in this cover type, often near a residual canopy tree. Hermit thrushes were less likely to inhabit mature deciduous forest, even-aged harvests, and harvested edges but chose nesting areas in mature mixed forest that was disturbed by road building and the seeding of landings and skid trails >10 years ago. Microhabitat characteristics of landcovers did not differ overall. Our results suggest a relationship with partial harvesting that is positive for wood thrush but negative for the other three species. ?? 2007 Elsevier B.V. All rights reserved.

Forest Ecology and Management

Atlantic Flyway review: Region V: Laurel, Prince Georges County, MD (390-0765)

Despite a 22% increase in net-hours, the number of new birds banded dropped 26% from 1977. The decrease was noted in each month and was spread among all of the commonest species, including summer residents, winter residents, permanent residents, and transients. The greatest declines were in Scarlet Tanager (84%), Blue Jay (78%), Red-eyed Vireo (67%), Gray Catbird (62%), Ruby-crowned Kinglet (60%}, Mrytle Warbler (56%), Gray-cheeked Thrush (55%), Wood Thrush (44%0, Cardinal (38%), and White-throated Sparrow (30%). Of these species, only the Scarlet Tanager and Myrtle Warbler had been banded in unusually large numbers in 1977. Partly offsetting the decreases were a few increases: Veery (from 4 to 22 birds), Black-and-white Warbler (3 to 12), and Ovenbird (15 to 34). The four species that had been singled out for mention last year as being victims of the severe winter weather in 1976-77 - the Winter and Carolina Wrens and the Golden- and Ruby-crowned Kinglets - fared no better in 1978. The first three were banded in almost identical nnmbers to 1977, and the Ruby-crown dropped again as indicated in the first paragraph.

Maryland

Modeling spatial variation in avian survival and residency probabilities

The importance of understanding spatial variation in processes driving animal population dynamics is widely recognized. Yet little attention has been paid to spatial modeling of vital rates. Here we describe a hierarchical spatial autoregressive model to provide spatially explicit year-specific estimates of apparent survival (phi) and residency (pi) probabilities from capture-recapture data. We apply the model to data collected on a declining bird species, Wood Thrush (Hylocichla mustelina), as part of a broad-scale bird-banding network, the Monitoring Avian Productivity and Survivorship (MAPS) program. The Wood Thrush analysis showed variability in both phi and pi among years and across space. Spatial heterogeneity in residency probability was particularly striking, suggesting the importance of understanding the role of transients in local populations. We found broad-scale spatial patterning in Wood Thrush phi and pi that lend insight into population trends and can direct conservation and research. The spatial model developed here represents a significant advance over approaches to investigating spatial pattern in vital rates that aggregate data at coarse spatial scales and do not explicitly incorporate spatial information in the model. Further development and application of hierarchical capture-recapture models offers the opportunity to more fully investigate spatiotemporal variation in the processes that drive population changes.

Ecology

Costs of detection bias in index-based population monitoring

Managers of wildlife populations commonly rely on indirect, count-based measures of the population in making decisions regarding conservation, harvest, or control. The main appeal in the use of such counts is their low material expense compared to methods that directly measure the population. However, their correct use rests on the rarely-tested but often-assumed premise that they proportionately reflect population size, i.e., that they constitute a population index. This study investigates forest management for the endangered Red-cockaded Woodpecker (Picoides borealis) and the Wood Thrush (Hylocichla mustelina) at the Piedmont National Wildlife Refuge in central Georgia, U.S.A. Optimal decision policies for a joint species objective were derived for two alternative models of Wood Thrush population dynamics. Policies were simulated under scenarios of unbiasedness, consistent negative bias, and habitat-dependent negative bias in observed Wood Thrush densities. Differences in simulation outcomes between biased and unbiased detection scenarios indicated the expected loss in resource objectives (here, forest habitat and birds) through decision-making based on biased population counts. Given the models and objective function used in our analysis, expected losses were as great as 11%, a degree of loss perhaps not trivial for applications such as endangered species management. Our analysis demonstrates that costs of uncertainty about the relationship between the population and its observation can be measured in units of the resource, costs which may offset apparent savings achieved by collecting uncorrected population counts.

Animal Biodiversity and Conservation

An integrated population model for bird monitoring in North America

Integrated population models (IPMs) provide a unified framework for simultaneously analyzing data sets of different types to estimate vital rates, population size, and dynamics; assess contributions of demographic parameters to population changes; and assess population viability. Strengths of an IPM include the ability to estimate latent parameters and improve the precision of parameter estimates. We present a hierarchical IPM that combines two broad‐scale avian monitoring data sets: count data from the North American Breeding Bird Survey (BBS) and capture–recapture data from the Monitoring Avian Productivity and Survivorship (MAPS) program. These data sets are characterized by large numbers of sample sites and observers, factors capable of inducing error in the sampling and observation processes. The IPM integrates the data sets by modeling the population abundance as a first‐order autoregressive function of the previous year's population abundance and vital rates. BBS counts were modeled as a log‐linear function of the annual index of population abundance, observation effects (observer identity and first survey year), and overdispersion. Vital rates modeled included adult apparent survival, estimated from a transient Cormack‐Jolly‐Seber model using MAPS data, and recruitment (surviving hatched birds from the previous season + dispersing adults) estimated as a latent parameter. An assessment of the IPM demonstrated it could recover true parameter values from 200 simulated data sets. The IPM was applied to data sets (1992–2008) of two bird species, Gray Catbird ( Dumetella carolinensis ) and Wood Thrush ( Hylocichla mustelina ) in the New England/Mid‐Atlantic coastal Bird Conservation Region of the United States. The Gray Catbird population was relatively stable (trend +0.4% per yr), while the Wood Thrush population nearly halved (trend −4.5% per yr) over the 17‐yr study period. IPM estimates of population growth rates, adult survival, and detection and residency probabilities were similar and as precise as estimates from the stand‐alone BBS and CJS models. A benefit of using the IPM was its ability to estimate the latent recruitment parameter. Annual growth rates for both species correlated more with recruitment than survival, and the relationship for Wood Thrush was stronger than for Gray Catbird. The IPM's unified modeling framework facilitates integration of these important data sets.

Ecological Applications

A productivity model for parasitized, multibrooded songbirds

We present an enhancement of a simulation model to predict annual productivity for Wood Thrushes (Hylocichla mustelina) and American Redstarts (Setophaga ruticilla); the model includes effects of Brown-headed Cowbird (Molothrus ater) parasitism. We used species-specific data from the Driftless Area Ecoregion of Wisconsin, Minnesota, and Iowa to parameterize the model as a case study. The simulation model predicted annual productivity of 2.03 ?? 1.60 SD for Wood Thrushes and 1.56 ?? 1.31 SD for American Redstarts. Our sensitivity analysis showed that high parasitism lowered Wood Thrush annual productivity more than American Redstart productivity, even though parasitism affected individual nests of redstarts more severely. Annual productivity predictions are valuable for habitat managers, but productivity is not easily obtained from field studies. Our model provides a useful means of integrating complex life history parameters to predict productivity for songbirds that experience nest parasitism. ?? The Cooper Ornithological Society 2006.

Condor

Use of radio-telemetry to reduce bias in nest searching

We used traditional searching, as well as radio-telemetry, to find 125 Wood Thrush ( Hylocichla mustelina ) nests during 1994–1996 at the Piedmont National Wildlife Refuge in Georgia, USA. We compared daily nest survival rates for 66 nests of radio-marked birds with 59 nests of birds found through systematic searching. By using radio-telemetry, we found Wood Thrush nests in higher elevation pine habitats, in addition to the more usual hardwood forests with moist soils. We found nests of radio-marked birds farther from streams than nests found by systematic searching. Thirty-two percent of radio-marked birds' nests were found at the tops of slopes, compared to 15% of the nests found by traditional searching. In addition, radio-marked birds generally moved up-slope for re-nesting attempts. Although the distribution of nests found with telemetry and searching varied, daily nest survival did not vary between the two groups. Radio-telemetry provided new information about Wood Thrush nesting habitats. We believe radio-telemetry can be a valuable addition to traditional searching techniques; it has the potential to provide a sample of nests free from a priori habitat biases.

Georgia

Thinking beyond the closure assumption: Designing surveys for estimating biological truth with occupancy models

Occupancy models estimate distributions of imperfectly detected species, but violations of the closure assumption can bias results. However, researchers working with mobile animals may find it impossible to eliminate such violations. Here, we tested the hypothesis that occupancy models fit to realistic sampling data can generate unbiased occupancy estimates for an itinerant Wood Thrush ( Hylocichla mustelina ) population. In 2013 and 2014, we tracked movements of 41 breeding Wood Thrush males. We modelled territory shift probabilities using logistic exposure models and within-territory movements using continuous-time stochastic process models. We then constructed an individual-based model, simulated (1000 iterations) spatiotemporal locations for individuals and simulated sampling these populations using 162 different point count protocols with variable spatial (sampling radius and point placement method), and temporal (survey length, between-survey intervals and number of surveys) characteristics. We compared occupancy estimates with true values of instantaneous, daily and seasonal occupancy from the simulations. We parameterized continuous time stochastic process models based on movements within 34 unique territories and estimated a daily territory shift probability of 0.0099 (95% CI: 0.0060, 0.0152). Simulated data indicated that estimates of occupancy ranged from 0.18 (0.06, 1.00) to 0.80 (0.71, 0.89) depending on protocol characteristics. Occupancy estimates increased with increasing survey radius, survey length and between-survey interval. Protocols using shorter surveys and between-survey intervals were good estimators for instantaneous occupancy (low bias and mean-squared error) but poor estimators for daily and seasonal occupancy; longer surveys and intervals generated unbiased estimators of daily occupancy but underestimated seasonal occupancy. Logistic regression models that ignored imperfect detection outperformed occupancy models for estimating instantaneous occupancy but not daily or seasonal occupancy. For mobile animals, occupancy of sampling sites changes in space and time. Consequently, the spatial and temporal aspects of a sampling protocol have strong, but predictable, effects on occupancy model parameter estimates. Our results demonstrate that how these factors interact is critical for designing surveys that produce occupancy estimates representative of the biological process of interest to a researcher.

Methods in Ecology and Evolution