Do the Golden-winged Warbler and Blue-winged Warbler Exhibit Species-specific Differences in their Breeding Habitat Use?

We compared habitat features of Golden-winged Warbler (Vermivora chrysoptera) territories in the presence and absence of the Blue-winged Warbler (V. cyanoptera) on reclaimed coal mines in southeastern Kentucky, USA. Our objective was to determine whether there are species specific differences in habitat that can be manipulated to encourage population persistence of the Golden-winged Warbler. When compared with Blue-winged Warblers, Golden-winged Warblers established territories at higher elevations and with greater percentages of grass and canopy cover. Mean territory size (minimum convex polygon) was 1.3 ha (se = 0.1) for Golden-winged Warbler in absence of Blue-winged Warbler, 1.7 ha (se = 0.3) for Golden-winged Warbler coexisting with Blue-winged Warbler, and 2.1 ha (se = 0.3) for Blue-winged Warbler. Territory overlap occurred within and between species (18 of n = 73 territories, 24.7%). All Golden-winged and Blue-winged Warblers established territories that included an edge between reclaimed mine land and mature forest, as opposed to establishing territories in open grassland/shrubland habitat. The mean distance territories extended from a forest edge was 28.0 m (se = 3.8) for Golden-winged Warbler in absence of Blue-winged Warbler, 44.7 m (se = 5.7) for Golden-winged Warbler coexisting with Blue-winged Warbler, and 33.1 m (se = 6.1) for Blue-winged Warbler. Neither territory size nor distances to forest edges differed significantly between Golden-winged Warbler in presence or absence of Blue-winged Warbler. According to Monte Carlo analyses, orchardgrass (Dactylis glomerata), green ash (Fraxinus pennsylvanica) seedlings and saplings, and black locust (Robinia pseudoacacia) saplings were indicative of sites with only Golden-winged Warblers. Sericea lespedeza, goldenrod (Solidago spp.), clematis vine (Clematis spp.), and blackberry (Rubus spp.) were indicative of sites where both species occurred. Our findings complement recent genetic studies and add another factor for examining Golden-winged Warbler population decline. Further, information from our study will aid land managers in manipulating habitat for the Golden-winged Warbler.

Modeling and Mapping Golden-winged Warbler Abundance to Improve Regional Conservation Strategies

Conservation planning requires identifying pertinent habitat factors and locating geographic locations where land management may improve habitat conditions for high priority species. I derived habitat models and mapped predicted abundance for the Golden-winged Warbler (Vermivora chrysoptera), a species of high conservation concern, using bird counts, environmental variables, and hierarchical models applied at multiple spatial scales. My aim was to understand habitat associations at multiple spatial scales and create a predictive abundance map for purposes of conservation planning for the Golden-winged Warbler. My models indicated a substantial influence of landscape conditions, including strong positive associations with total forest composition within the landscape. However, many of the associations I observed were counter to reported associations at finer spatial extents; for instance, I found Golden-winged Warblers negatively associated with several measures of edge habitat. No single spatial scale dominated, indicating that this species is responding to factors at multiple spatial scales. I found Golden-winged Warbler abundance was negatively related with Blue-winged Warbler (Vermivora cyanoptera) abundance. I also observed a north-south spatial trend suggestive of a regional climate effect that was not previously noted for this species. The map of predicted abundance indicated a large area of concentrated abundance in west-central Wisconsin, with smaller areas of high abundance along the northern periphery of the Prairie Hardwood Transition. This map of predicted abundance compared favorably with independent evaluation data sets and can thus be used to inform regional planning efforts devoted to conserving this species.

Use of Large Clear-Cuts by Wilson’s Warbler in an Eastern Canadian Boreal Forest

Wilson’s Warbler (Cardellina pusilla; WIWA) has been declining for several decades, possibly because of habitat loss. We compared occupancy of territorial males in two habitat types of Québec’s boreal forest, alder (Alnus spp.) scrubland and recent clear-cuts. Singing males occurred in clusters, their occupancy was similar in both habitats, but increased with the amount of alder or clear-cut within 400 m of point-count stations. A despotic distribution of males between habitats appeared unlikely, because there were no differences in morphology between males captured in clear-cuts vs. alder. Those results contrast with the prevailing view, mostly based on western populations, that WIWA are wetland or riparian specialists, and provide the first evidence for a preference for large tracts of habitat in this species. Clear-cuts in the boreal forest may benefit WIWA by supplying alternative nesting habitat. However, the role of clear-cuts as source or sink habitats needs to be addressed with data on reproduction.

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.

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.

Regional habitat needs of a nationally listed species, Canada Warbler (Cardellina canadensis), in Alberta, Canada

Understanding factors that affect the distribution and abundance of species is critical to developing effective management plans for conservation. Our goal was to quantify the distribution and abundance of Canada Warbler (Cardellina canadensis), a threatened old-forest associate in Alberta, Canada. The Canada Warbler has declined across its range, including in Alberta where habitat loss and alteration from urban expansion, forestry, and energy development are changing the forest landscape. We used 110,427 point count survey visits from 32,287 unique survey stations to model local-level (150-m radius circular buffers) and stand-level (564-m radius circular buffers) habitat associations of the Canada Warbler. We found that habitat supporting higher densities of Canada Warblers was locally concentrated yet broadly distributed across Alberta’s boreal forest region. Canada Warblers were most commonly associated with older deciduous forest at the local scale, particularly near small, incised streams, and greater amounts of deciduous forest at the stand scale. Predicted density was lower in other forest types and younger age classes measured at the local scale. There was little evidence that local-scale fragmentation (i.e., edges created by linear features) influenced Canada Warbler abundance. However, current forestry practices in the province likely will reduce the availability of Canada Warbler habitat over time by cutting old deciduous forest stands. Our results suggest that conservation efforts aimed at Canada Warbler focus on retaining large stands of old deciduous forest, specifically stands adjacent to streams, by increasing the width of deciduous retention buffers along streams during harvest and increasing the size and number of old forest residual patches in harvested stands.

Identification of Summer Origins of Songbirds Migrating through Southern Canada in Autumn

Stable hydrogen isotopes (δD) in flight feathers were measured to investigate the summer origins of five species of boreal-breeding warblers captured during fall migration at Canadian Migration Monitoring Network (CMMN) stations spread across southern Canada. Mean δD varied among stations and species within stations, but there was broad overlap in δD values. Although isotope ratios indicate that migrants at each station come from a wide range of latitudes, they are unable to provide much longitudinal discrimination. Band recoveries are sparse, but indicate that in general western Canadian warblers move southeast in fall, eastern birds move southwest, and there is a transition zone in the Great Lakes region. Combining knowledge of migratory direction with isotope results increases discrimination of breeding areas. Isotope results support fall migratory movements by Blackpoll Warbler (Dendroica striata) and Northern Waterthrush (Seiurus novaboracensis) that are more easterly than for other species, and in all study species, birds from more northern regions passed through southern Canada later in the season. Migration monitoring stations capture birds from broad areas of latitude, and migrants passing through each province appear to come from largely different portions of the Canadian breeding range, so a few stations placed in each province should suffice collectively to sample birds from most of the boreal forest. Migration monitoring in southern Canada, therefore, has the potential to monitor status of boreal forest birds in Canada that are unsampled by other monitoring programs.

Effects of Spruce Budworm (Choristoneura fumiferana (Clem.)) Outbreaks on Boreal Mixed-Wood Bird Communities

This study examined the influence of a spruce budworm (Choristoneura fumiferana (Clem.)) outbreak on a boreal mixed-wood bird community in forest stands ranging in age from 0 to 223 yr. We asked if (1) patterns of species response were consistent with the existence of spruce budworm specialists, i.e., species that respond in a stronger quantitative or qualitative way than other species; (2) the superabundance of food made it possible for species to expand their habitat use in age classes that were normally less used; and (3) the response to budworm was limited to specialists or was it more widespread. Results here indicated that three species, specifically the Bay-breasted Warbler (Dendroica castanea), Tennessee Warbler (Vermivora peregrina), and Cape May Warbler (Dendroica tigrina), had a larger numerical response to the budworm outbreak. They responded with increases in density of up to tenfold over 4 or 5 yr. No other species responded with more than a twofold increase in the same time period. These species also showed a functional response by breeding more frequently in young stands aged 1–21 yr and intermediate stands aged 22–36 yr as budworm numbers increased. Our data also suggested that many species profited to a lesser extent from budworm outbreaks, but that this effect may be too subtle to detect in most studies. We found evidence of a positive numerical effect in at least 18 additional species in one or two stand-age categories but never in all three for any one species. Given the numerical response in many species and the potential influence of budworm on bird populations because of the vast extent of outbreaks, we believe that the population cycle of spruce budworm should be considered in any evaluation of population trends in eastern boreal birds.

Contrasting Seasonal Survivorship of Two Migratory Songbirds Wintering in Threatened Mangrove Forests

Long-distance migrants wintering in tropical regions face a number of critical conservation threats throughout their lives, but seasonal estimates of key demographic parameters such as winter survival are rare. Using mist-netting-based mark-recapture data collected in coastal Costa Rica over a six-year period, we examined variation in within- and between-winter survivorship of the Prothonotary Warbler (Protonotaria citrea; 753 young and 376 adults banded), a declining neotropical habitat specialist that depends on threatened mangrove forests during the nonbreeding season. We derived parallel seasonal survivorship estimates for the Northern Waterthrush (Seiurus noveboracensis; 564 young and 93 adults banded), a cohabitant mangrove specialist that has not shown the same population decline in North America, to assess whether contrasting survivorship might contribute to the observed differences in the species’ population trajectories. Although average annual survival probability was relatively similar between the two species for both young and adult birds, monthly estimates indicated that relative to Northern Waterthrush, Prothonotary Warblers exhibited: greater interannual variation in survivorship, especially within winters; greater variation in survivorship among the three study sites; lower average between-winter survivorship, particularly among females, and; a sharp decline in between-winter survivorship from 2003 to 2009 for both age groups and both sexes. Rather than identifying one seasonal vital rate as a causal factor of Prothonotary Warbler population declines, our species comparison suggests that the combination of variable within-winter survival with decreasing between-winter survival demands a multi-seasonal approach to the conservation of this and other tropical-wintering migrants.

Numerical Response of Breeding Birds Following Experimental Selection Harvesting in Northern Hardwood Forests

Silvicultural treatments have been shown to alter the composition of species assemblages in numerous taxa. However, the intensity and persistence of these effects have rarely been documented. We used a before-after, control-impact (BACI) paired design, i.e., five pairs of 25-ha study plots, 1-control and 1-treated plot, to quantify changes in the density of eight forest bird species in response to selection harvesting over six breeding seasons, one year pre- and five years postharvest. Focal species included mature forest associates, i.e., Northern Parula (Setophaga americana) and Black-throated Green Warbler (Setophaga virens), forest generalists, i.e., Yellow-bellied Sapsucker (Sphyrapicus varius) and Swainson’s Thrush (Catharus ustulatus), early-seral specialists, i.e., Mourning Warbler (Geothlypis philadelphia) and Chestnut-sided Warbler (Setophaga pensylvanica), species associated with shrubby forest gaps, i.e., Black-throated Blue Warbler (Setophaga caerulescens), and mid-seral species, i.e., American Redstart (Setophaga ruticilla). As predicted, we found a negative numerical response to the treatment in the Black-throated Green Warbler, no treatment effect in the Yellow-bellied Sapsucker, and a positive treatment effect in early-seral specialists. We only detected a year effect in the Northern Parula and the American Redstart. There was evidence for a positive treatment effect on the Swainson’s Thrush when the regeneration started to reach the pole stage, i.e., fifth year postharvest. These findings suggest that selection harvesting has the potential to maintain diverse avian assemblages while allowing sustainable management of timber supply, but future studies should determine whether mature-forest associates can sustain second- and third-entry selection harvest treatments.

Age-specific survival of male Golden-cheeked Warblers on the Fort Hood Military Reservation, Texas

Population models are essential components of large-scale conservation and management plans for the federally endangered Golden-cheeked Warbler (Setophaga chrysoparia; hereafter GCWA). However, existing models are based on vital rate estimates calculated using relatively small data sets that are now more than a decade old. We estimated more current, precise adult and juvenile apparent survival (Φ) probabilities and their associated variances for male GCWAs. In addition to providing estimates for use in population modeling, we tested hypotheses about spatial and temporal variation in Φ. We assessed whether a linear trend in Φ or a change in the overall mean Φ corresponded to an observed increase in GCWA abundance during 1992-2000 and if Φ varied among study plots. To accomplish these objectives, we analyzed long-term GCWA capture-resight data from 1992 through 2011, collected across seven study plots on the Fort Hood Military Reservation using a Cormack-Jolly-Seber model structure within program MARK. We also estimated Φ process and sampling variances using a variance-components approach. Our results did not provide evidence of site-specific variation in adult Φ on the installation. Because of a lack of data, we could not assess whether juvenile Φ varied spatially. We did not detect a strong temporal association between GCWA abundance and Φ. Mean estimates of Φ for adult and juvenile male GCWAs for all years analyzed were 0.47 with a process variance of 0.0120 and a sampling variance of 0.0113 and 0.28 with a process variance of 0.0076 and a sampling variance of 0.0149, respectively. Although juvenile Φ did not differ greatly from previous estimates, our adult Φ estimate suggests previous GCWA population models were overly optimistic with respect to adult survival. These updated Φ probabilities and their associated variances will be incorporated into new population models to assist with GCWA conservation decision making.

Variables associated with nest survival of Golden-winged Warblers (Vermivora chrysoptera) among vegetation communities commonly used for nesting

Among shrubland- and young forest-nesting bird species in North America, Golden-winged Warblers (Vermivora chrysoptera) are one of the most rapidly declining partly because of limited nesting habitat. Creation and management of high quality vegetation communities used for nesting are needed to reduce declines. Thus, we examined whether common characteristics could be managed across much of the Golden-winged Warbler’s breeding range to increase daily survival rate (DSR) of nests. We monitored 388 nests on 62 sites throughout Minnesota, Wisconsin, New York, North Carolina, Pennsylvania, Tennessee, and West Virginia. We evaluated competing DSR models in spatial-temporal (dominant vegetation type, population segment, state, and year), intraseasonal (nest stage and time-within-season), and vegetation model suites. The best-supported DSR models among the three model suites suggested potential associations between daily survival rate of nests and state, time-within-season, percent grass and Rubus cover within 1 m of the nest, and distance to later successional forest edge. Overall, grass cover (negative association with DSR above 50%) and Rubus cover (DSR lowest at about 30%) within 1 m of the nest and distance to later successional forest edge (negative association with DSR) may represent common management targets across our states for increasing Golden-winged Warbler DSR, particularly in the Appalachian Mountains population segment. Context-specific adjustments to management strategies, such as in wetlands or areas of overlap with Blue-winged Warblers (Vermivora cyanoptera), may be necessary to increase DSR for Golden-winged Warblers.