Spatial Aggregation of Forest Songbird Territories and Possible Implications for Area Sensitivity

Habitat area requirements of forest songbirds vary greatly among species, but the causes of this variation are not well understood. Large area requirements could result from advantages for certain species when settling their territories near those of conspecifics. This phenomenon would result in spatial aggregations much larger than single territories. Species that aggregate their territories could show reduced population viability in highly fragmented forests, since remnant patches may remain unoccupied if they are too small to accommodate several territories. The objectives of this study were twofold: (1) to seek evidence of territory clusters of forest birds at various spatial scales, lags of 250-550 m, before and after controlling for habitat spatial patterns; and (2) to measure the relationship between spatial autocorrelation and apparent landscape sensitivity for these species. In analyses that ignored spatial variation of vegetation within remnant forest patches, nine of the 17 species studied significantly aggregated their territories within patches. After controlling for forest vegetation, the locations of eight out of 17 species remained significantly clustered. The aggregative pattern that we observed may, thus, be indicative of a widespread phenomenon in songbird populations. Furthermore, there was a tendency for species associated with higher forest cover to be more spatially aggregated [ERRATUM].

Forest Policy Scenario Analysis: Sensitivity of Songbird Community to Changes in Forest Cover Amount and Configuration

Changes in mature forest cover amount, composition, and configuration can be of significant consequence to wildlife populations. The response of wildlife to forest patterns is of concern to forest managers because it lies at the heart of such competing approaches to forest planning as aggregated vs. dispersed harvest block layouts. In this study, we developed a species assessment framework to evaluate the outcomes of forest management scenarios on biodiversity conservation objectives. Scenarios were assessed in the context of a broad range of forest structures and patterns that would be expected to occur under natural disturbance and succession processes. Spatial habitat models were used to predict the effects of varying degrees of mature forest cover amount, composition, and configuration on habitat occupancy for a set of 13 focal songbird species. We used a spatially explicit harvest scheduling program to model forest management options and simulate future forest conditions resulting from alternative forest management scenarios, and used a process-based fire-simulation model to simulate future forest conditions resulting from natural wildfire disturbance. Spatial pattern signatures were derived for both habitat occupancy and forest conditions, and these were placed in the context of the simulated range of natural variation. Strategic policy analyses were set in the context of current Ontario forest management policies. This included use of sequential time-restricted harvest blocks (created for Woodland caribou (Rangifer tarandus) conservation) and delayed harvest areas (created for American marten (Martes americana atrata) conservation). This approach increased the realism of the analysis, but reduced the generality of interpretations. We found that forest management options that create linear strips of old forest deviate the most from simulated natural patterns, and had the greatest negative effects on habitat occupancy, whereas policy options that specify deferment and timing of harvest for large blocks helped ensure the stable presence of an intact mature forest matrix over time. The management scenario that focused on maintaining compositional targets best supported biodiversity objectives by providing the composition patterns required by the 13 focal species, but this scenario may be improved by adding some broad-scale spatial objectives to better maintain large blocks of interior forest habitat through time.

The Importance of Agriculture-Dominated Landscapes and Lack of Field Border Effect for Early-Succession Songbird Nest Success

In recent decades, many early-succession songbird species have experienced severe and widespread declines, which often are related to habitat destruction. Field borders create additional or enhance existing early-succession habitat on farmland. However, field border shape and the landscape context surrounding farms may influence the effectiveness of field borders in contributing to the stabilization or increase of early-succession bird populations. We examined the influence of linear and nonlinear field borders on farms in landscapes dominated by either agriculture or forests on nest success and Brown-headed Cowbird (Molothrus ater) brood parasitism of Indigo Bunting (Passerina cyanea) and Blue Grosbeak (Passerina caerulea) nests combined. Field border establishment did not affect nest survival probability and brood parasitism frequency of Indigo Bunting and Blue Grosbeak nests. Indigo Bunting/Blue Grosbeak nest success probability was more than twice as high in agriculture-dominated landscapes (39%) than in forested landscapes (17%), and brood parasitism frequency was high (33%) but did not differ between landscapes. Edges in agriculture-dominated landscapes can be higher-quality habitats for early-succession birds than edges in forest-dominated landscapes, but our field border treatments did not enhance nest success for these birds on farms in either landscape.

How well do regional or national Breeding Bird Survey data predict songbird population trends at an intact boreal site?

A study to monitor boreal songbird trends was initiated in 1998 in a relatively undisturbed and remote part of the boreal forest in the Northwest Territories, Canada. Eight years of point count data were collected over the 14 years of the study, 1998-2011. Trends were estimated for 50 bird species using generalized linear mixed-effects models, with random effects to account for temporal (repeat sampling within years) and spatial (stations within stands) autocorrelation and variability associated with multiple observers. We tested whether regional and national Breeding Bird Survey (BBS) trends could, on average, predict trends in our study area. Significant increases in our study area outnumbered decreases by 12 species to 6, an opposite pattern compared to Alberta (6 versus 15, respectively) and Canada (9 versus 20). Twenty-two species with relatively precise trend estimates (precision to detect > 30% decline in 10 years; observed SE ≤ 3.7%/year) showed nonsignificant trends, similar to Alberta (24) and Canada (20). Precision-weighted trends for a sample of 19 species with both reliable trends at our site and small portions of their range covered by BBS in Canada were, on average, more negative for Alberta (1.34% per year lower) and for Canada (1.15% per year lower) relative to Fort Liard, though 95% credible intervals still contained zero. We suggest that part of the differences could be attributable to local resource pulses (insect outbreak). However, we also suggest that the tendency for BBS route coverage to disproportionately sample more southerly, developed areas in the boreal forest could result in BBS trends that are not representative of range-wide trends for species whose range is centred farther north.

Factors driving territory size and breeding success in a threatened migratory songbird, the Canada Warbler

Successful conservation of migratory birds demands we understand how habitat factors on the breeding grounds influences breeding success. Multiple factors are known to directly influence breeding success in territorial songbirds. For example, greater food availability and fewer predators can have direct effects on breeding success. However, many of these same habitat factors can also result in higher conspecific density that may ultimately reduce breeding success through density dependence. In this case, there is a negative indirect effect of habitat on breeding success through its effects on conspecific density and territory size. Therefore, a key uncertainty facing land managers is whether important habitat attributes directly influence breeding success or indirectly influence breeding success through territory size. We used radio-telemetry, point-counts, vegetation sampling, predator observations, and insect sampling over two years to provide data on habitat selection of a steeply declining songbird species, the Canada Warbler (Cardellina canadensis). These data were then applied in a hierarchical path modeling framework and an AIC model selection approach to determine the habitat attributes that best predict breeding success. Canada Warblers had smaller territories in areas with high shrub cover, in the presence of red squirrels (Tamiasciurus hudsonicus), at shoreline sites relative to forest-interior sites and as conspecific density increased. Breeding success was lower for birds with smaller territories, which suggests competition for limited food resources, but there was no direct evidence that food availability influenced territory size or breeding success. The negative relationship between shrub cover and territory size in our study may arise because these specific habitat conditions are spatially heterogeneous, whereby individuals pack into patches of preferred breeding habitat scattered throughout the landscape, resulting in reduced territory size and an associated reduction in resource availability per territory. Our results therefore highlight the importance of considering direct and indirect effects for Canada warblers; efforts to increase the amount of breeding habitat may ultimately result in lower breeding success if habitat availability is limited and negative density dependent effects occur.

Understanding Demographic and Behavioral Mechanisms that Guide Responses of Neotropical Migratory Birds to Urbanization: a Simulation Approach

Although studies often report that densities of many forest birds are negatively related to urbanization, the mechanisms guiding this pattern are poorly understood. Our objective was to use a population simulation to examine the relative influence of six demographic and behavioral processes on patterns of avian abundance in urbanizing landscapes. We constructed an individual-based population simulation model representing the annual cycle of a Neotropical migratory songbird. Each simulation was performed under two landscape scenarios. The first scenario had similar proportions of high- and low-quality habitat across the urban to rural gradient. Under the first scenario, avian density was negatively related to urbanization only when rural habitats were perceived to be of higher quality than they actually were. The second landscape scenario had declining proportions of high-quality habitat as urbanization increased. Under the second scenario, each mechanism generated a negative relationship between density and urbanization. The strongest effect on density resulted when birds preferentially selected habitats in landscapes from which they fledged or were constrained from dispersing. The next strongest patterns occurred when birds directly evaluated habitat quality and accurately selected the highest-quality available territories. When birds selected habitats based on the presence of conspecifics, the density–urbanization relationship was only one-third the strength of other habitat selection mechanisms and only occurred under certain levels of population survival. Although differences in adult or nest survival in the face of random habitat selection still elicited reduced densities in urban landscapes, the relationships between urbanization and density were weaker than those produced by the conspecific attraction mechanism. Results from our study identify key predictions and areas for future research, including assessing habitat quality in urban and rural areas in order to determine if habitats in urban areas are underutilized.

Movement of Juvenile Songbirds in Harvested Boreal Forest: Assessing Residency Time and Landscape Connectivity

Little is known about juvenile songbird movement in response to timber harvest, particularly in the boreal forest. If clearcut land cover facilitates movement, the availability of resources may increase. However, if clearcut land cover impedes movement, important post-fledging resources may be rendered inaccessible. Using radio telemetry, we tested the hypothesis that regenerating clearcut land cover would affect the movement of recently independent Yellow-rumped Myrtle Warblers (Dendroica coronata coronata) and Blackpoll Warblers (Dendroica striata) differently than forested land cover owing to intrinsic differences in each land-cover type or in how they are perceived. We found that both species moved extensively before migration. We also found that Blackpoll Warblers were quick to exit local areas composed of clearcut land cover and that both species were quick to exit neighborhoods composed of large proportions of clearcut land cover. However, if individuals encountered clearcut land cover when exiting the neighborhood, movement rate was slowed. Effectively, residency time decreased in clearcut neighborhoods and landscape connectivity was impeded by clearcut land cover. Our results suggest that clearcut land cover may represent low-quality habitat for both species during the post-fledging period. Further research is needed to determine if changes in movement behavior associated with landscape structure affect individual condition and higher-level ecological processes.

Ovenbird (Seiurus aurocapilla) Territory Placement Near Seismic Lines is Influenced by Forest Regeneration and Conspecific Density

The boreal forest of western Canada is being dissected by seismic lines used for oil and gas exploration. The vast amount of edge being created is leading to concerns that core habitat will be reduced for forest interior species for extended periods of time. The Ovenbird (Seiurus aurocapilla) is a boreal songbird known to be sensitive to newly created seismic lines because it does not include newly cut lines within its territory. We examined multiple hypotheses to explain potential mechanisms causing this behavior by mapping Ovenbird territories near lines with varying states of vegetation regeneration. The best model to explain line exclusion behavior included the number of neighboring conspecifics, the amount of bare ground, leaf-litter depth, and canopy closure. Ovenbirds exclude recently cut seismic lines from their territories because of lack of protective cover (lower tree and shrub cover) and because of reduced food resources due to large areas of bare ground. Food reduction and perceived predation risk effects seem to be mitigated once leaf litter (depth and extent of cover) and woody vegetation cover are restored to forest interior levels. However, as conspecific density increases, lines are more likely to be used as landmarks to demarcate territorial boundaries, even when woody vegetation cover and leaf litter are restored. This behavior can reduce territory density near seismic lines by changing the spatial distribution of territories. Landmark effects are longer lasting than the effects from reduced food or perceived predation risk because canopy height and tree density take >40 years to recover to forest interior levels. Mitigation of seismic line impacts on Ovenbirds should focus on restoring forest cover as quickly as possible after line cutting.

Crop type influences edge effects on the reproduction of songbirds in sagebrush habitat near agriculture

Extensive fragmentation of the sagebrush shrubsteppe of western North America could be contributing to observed population declines of songbirds in sagebrush habitat. We examined whether habitat fragmentation impacts the reproduction of songbirds in sagebrush edge habitat near agriculture, and if potential impacts vary depending on the adjacent crop type. Specifically, we evaluated whether nest abundance and nest survival varied between orchard edge habitat, vineyard edge habitat, and interior habitat. We then examined whether the local nest predator community and vegetation could explain the differences detected. We detected fewer nests in edge than interior habitat. Nest abundance per songbird was also lower in edge than interior habitat, although only adjacent to vineyards. Nest predation was more frequent in orchard edge habitat than vineyard edge or interior habitat. Predators identified with nest cameras were primarily snakes, however, reduced nest survival in orchard edge habitat was not explained by differences in the abundance of snakes or any other predator species identified. Information theoretic analysis of daily survival rates showed that greater study plot shrub cover and lower grass height at nests were partially responsible for the lower rate of predation-specific daily nest survival rate (PDSR) observed in orchard edge habitat, but additional factors are likely important. Results of this study suggest that different crop types have different edge effects on songbirds nesting in sagebrush shrubsteppe, and that these reproductive edge effects may contribute to observed declines of these species. Habitat managers should avoid the creation of new orchard-sagebrush habitat edges to avoid further impacts on already declining songbird populations.

Stand and within-stand factors influencing Golden-winged Warbler use of regenerating stands in the central Appalachian Mountains

The Golden-winged Warbler (Vermivora chrysoptera) is currently being considered for protected status under the U.S. Endangered Species Act. The creation of breeding habitat in the Appalachian Mountains is considered a conservation priority for this songbird, which is dependent on extensively forested landscapes with adequate availability of young forest. We modeled abundance of Golden-winged Warbler males in regenerating harvested forest stands that were 0-17 years postharvest at both mid-Appalachian and northeast Pennsylvania regional scales using stand and within-stand characteristics of 222 regenerating stands, 2010-2011. Variables that were most influential at the mid-Appalachian scale were different than those in the northeast region. Across the mid-Appalachian ecoregion, the proportion of young forest cover, i.e., shrub/scrub cover, within 1 km of regenerating stands best explained abundance of Golden-winged Warblers. Golden-winged Warbler response was best explained by a concave quadratic relationship in which abundance was highest with 5-15% land in young forest cover. We also found evidence that the amount of herbaceous cover, i.e., the amount of grasses and forbs, within a regenerating stand positively influenced abundance of Golden-winged Warblers. In northeastern Pennsylvania, where young forest cover is found in high proportions, the distance to the nearest regenerating stand best explained variation in abundance of Golden-winged Warblers. Abundance of Golden-winged Warblers was <1 male per survey when another regenerating stand was >1500 m away. When modeling within-stand features in the northeast region, many of the models were closely ranked, indicating that multiple variables likely explained Golden-winged Warbler response to within-stand conditions. Based on our findings, we have proposed several management guidelines for land managers interested in creating breeding habitat for Golden-winged Warblers using commercial timber operations. For example, we recommend when managing for Golden-winged Warblers in the central Appalachian Mountains that managers should strive for 15% young forest in a heavily forested landscape (>70% forest cover) and cluster stands within 1-2 km of other young forest habitats.

Effects of Gunnison Sage-Grouse habitat treatment efforts on associated avifauna and vegetation structure

Conservation efforts over the last 20 years for the Gunnison Sage-Grouse (Centrocercus minimus) have involved extensive habitat manipulations done predominantly to improve brood rearing habitat for the grouse. However, the effects of Gunnison Sage-Grouse habitat treatments on sympatric avifauna and responses of vegetation to manipulations are rarely measured, and if they are, it is immediately following treatment implementation. This study examined the concept of umbrella species management by retrospectively comparing density and occupancy of eight sagebrush associated songbird species and six measures of vegetation in treated and control sites. Our results suggested that songbird densities and occupancy changed for birds at the extreme ends of their association with sagebrush and varied with fine-scale habitat structure. We found Brewer’s Sparrows (Spizella breweri) decreased in density on treated sites and Vesper Sparrows (Pooecetes gramineus) increased. Occupancy estimation revealed that Brewer’s Sparrows and Green-tailed Towhees (Pipilo chlorurus) occupied significantly fewer treated points whereas Vesper Sparrows occupied significantly more. Vegetation comparisons between treated and control areas found shrub cover to be 26% lower in treated sites. Lower shrub cover in treated areas may explain the differences in occupancy and densities of the species sampled based on known habitat needs. The fine-scale analysis showed a negative relationship to forb height and cover for the Sage Sparrow (Amphispiza belli) indicating, from vegetation measures showing grass and forb cover during a good precipitation year covered significantly more area in the treatment than the control sites, that Sage Sparrows may also not respond favorably to Gunnison Sage-Grouse habitat treatments. While the concept of an umbrella species is appealing, evidence from this study suggests that conservation efforts aimed at the Gunnison Sage-Grouse may not be particularly effective for conserving other sagebrush obligate species of concern. This is probably due to Gunnison Sage-Grouse habitat management being focused on the improvement of brood rearing habitat which reduces sagebrush cover and promotes development of understory forbs and grasses.