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.

California's Endangered Belding’s Savannah Sparrow (Passerculus sandwichensis beldingi): Tolerance of Pedestrian Disturbance

Salt marshes constitute habitat islands for many endemic animal species, particularly along the California coast, where urban sprawl has fragmented this habitat. Recreational activities in salt marshes have increased recently, posing an interesting problem: how do endemic species lacking alternative habitat modify their tolerance to humans? We assessed seasonal and site variations in three tolerance parameters (distances at which animals became alert, fled, and moved after fleeing) of California's endangered Belding’s Savannah Sparrow ((Passerculus sandwichensis beldingi). We approached individuals on trails in three salt marshes with different levels of vehicle and pedestrian traffic. Belding’s Savannah Sparrows became aware and fled at shorter distances in the salt marsh coincident with greater levels of recreational activity as a result of habituation or visual obstruction effects. Seasonal effects in tolerance varied between sites. Alert and flight initiation distances were higher in the pre-nesting than in the non-breeding season in the site with the highest levels of recreational use likely due to greater exposure of breeding individuals; however, the opposite seasonal trend was found in each of the two sites with relatively lower human use, probably because individuals were less spatially attached in the non-breeding season when they foraged in aggregations. Distance fled was greater in the non-breeding than in the breeding season. Our findings call for dynamic management of recreational activities in different salt marshes depending on the degree of exposure to humans and seasonal variations in tolerance. We recommend a minimum approaching distance of 63 m and buffer areas of 1.3 ha around Belding's Savannah Sparrows.

Linking Canadian Harvested Juvenile American Black Ducks to Their Natal Areas Using Stable Isotope (δD, δ13C, and δ15N) Methods

Understanding source-sink dynamics of game birds is essential to harvest and habitat management but acquiring this information is often logistically and financially challenging using traditional methods of population surveys and banding studies. This is especially true for species such as the American Black Duck (Anas rubripes), which have low breeding densities and extensive breeding ranges that necessitate extensive surveys and banding programs across eastern North America. Despite this effort, the contribution of birds fledged from various landscapes and habitat types within specific breeding ranges to regional harvest is largely unknown but remains an important consideration in adaptive harvest management and targeted habitat conservation strategies. We investigated if stable isotope (δD, δ13C, δ15N) could augment our present understanding of connectivity between breeding and harvest areas and so provide information relevant to the two main management strategies for black ducks, harvest and habitat management. We obtained specimens from 200 hatch-year Black Duck wings submitted to the Canadian Wildlife Service Species Composition Survey. Samples were obtained from birds harvested in Western, Central, and Eastern breeding/harvest subregions to provide a sample representative of the range and harvest rate of birds harvested in Canada. We sampled only hatch-year birds to provide an unambiguous and direct link between production and harvest areas. Marine origins were assigned to 12%, 7%, and 5% of birds harvested in the Eastern, Central, and Western subregions, respectively. In contrast, 32%, 9%, and 5% of birds were assigned, respectively, to agricultural origins. All remaining birds were assigned to nonagricultural origins. We portrayed probability of origin using a combination of Bayesian statistical and GIS methods. Placement of most eastern birds was western Nova Scotia, eastern New Brunswick, Prince Edward Island, and southern Newfoundland. Agricultural birds from the Central region were consistent with the Saguenay region of Québec and the eastern claybelt with nonagricultural birds originating in the boreal. Western nonagricultural birds were associated with broad boreal origins from southern James Bay to Lake of the Woods and east to Cochrane, Ontario. Our work shows that the geographic origins, landscape, and habitat associations of hatch-year Black Ducks can be inferred using this technique and we recommend that a broad-scale isotopic study using a large sample of Canadian and US harvested birds be implemented to provide a continental perspective of source-sink population dynamics.

Reproductive Consequences of Nest Site Use in Fork-tailed Storm-Petrels in the Aleutian Islands, Alaska: Potential Lasting Effects of an Introduced Predator

We examined the reproductive consequences of differential nest site use in Fork-tailed Storm-Petrels (Oceanodroma furcata) in the Aleutian Islands, Alaska, where birds on islands where foxes were introduced nest in rocky substrate rather than in typical soil habitat. We investigated how physical and microclimatic nest site characteristics influenced storm-petrel breeding success 20 years after fox removal. We then examined whether those nest site characteristics that affected success were related to the amount of rock that composed the nest. In both years of our study, nest temperature had the strongest influence on chick survival and overall reproductive success, appearing in all the top models and alone explaining 14–35% of the variation in chick survival. The relationship between reproductive success and nest temperature was positive in both years, with higher survival in warmer nests. In turn, the best predictor of nest temperature was the amount of rock that composed the site. Rockier nests had colder average temperatures, which were driven by lower daily minimum temperatures, compared to nests with more soil. Thus, the rockiness of the nest site appeared to affect chick survival and overall reproductive success through its influence on nest temperature. This study suggests that the use of rocky nest sites, presumed to be a result of historic predation from introduced foxes, could decrease breeding success in this recovering population, and thus be a long-lasting effect of introduced predators.

Identification of Putative Wintering Areas and Ecological Determinants of Population Dynamics of Common House-Martin (Delichon urbicum) and Common Swift (Apus apus) Breeding in Northern Italy

To identify the causes of population decline in migratory birds, researchers must determine the relative influence of environmental changes on population dynamics while the birds are on breeding grounds, wintering grounds, and en route between the two. This is problematic when the wintering areas of specific populations are unknown. Here, we first identified the putative wintering areas of Common House-Martin (Delichon urbicum) and Common Swift (Apus apus) populations breeding in northern Italy as those areas, within the wintering ranges of these species, where the winter Normalized Difference Vegetation Index (NDVI), which may affect winter survival, best predicted annual variation in population indices observed in the breeding grounds in 1992–2009. In these analyses, we controlled for the potentially confounding effects of rainfall in the breeding grounds during the previous year, which may affect reproductive success; the North Atlantic Oscillation Index (NAO), which may account for climatic conditions faced by birds during migration; and the linear and squared term of year, which account for nonlinear population trends. The areas thus identified ranged from Guinea to Nigeria for the Common House-Martin, and were located in southern Ghana for the Common Swift. We then regressed annual population indices on mean NDVI values in the putative wintering areas and on the other variables, and used Bayesian model averaging (BMA) and hierarchical partitioning (HP) of variance to assess their relative contribution to population dynamics. We re-ran all the analyses using NDVI values at different spatial scales, and consistently found that our population of Common House-Martin was primarily affected by spring rainfall (43%–47.7% explained variance) and NDVI (24%–26.9%), while the Common Swift population was primarily affected by the NDVI (22.7%–34.8%). Although these results must be further validated, currently they are the only hypotheses about the wintering grounds of the Italian populations of these species, as no Common House-Martin and Common Swift ringed in Italy have been recovered in their wintering ranges.

Risk of Agricultural Practices and Habitat Change to Farmland Birds

Many common bird species have declined as a result of agricultural intensification and this could be mitigated by organic farming. We paired sites for habitat and geographical location on organic and nonorganic farms in Ontario, Canada to test a priori predictions of effects on birds overall, 9 guilds and 22 species in relation to candidate models for farming practices (13 variables), local habitat features (12 variables), or habitat features that influence susceptibility to predation. We found that: (1) Overall bird abundance, but not richness, was significantly (p < 0.05) higher on organic sites (mean 43.1 individuals per site) than nonorganic sites (35.8 individuals per site). Significantly more species of birds were observed for five guilds, including primary grassland birds, on organic vs. nonorganic sites. No guild had higher richness or abundance on nonorganic farms; (2) Farming practice models were the best (ΔAIC < 4) for abundance of birds overall, primary grassland bird richness, sallier aerial insectivore richness and abundance, and abundance of ground nesters; (3) Habitat models were the best for overall richness, Neotropical migrant abundance, richness and abundance of Ontario-USA-Mexico (short-distance) migrants and resident richness; (4) Predation models were the best for richness of secondary grassland birds and ground feeders; (5) A combination of variables from the model types were best for richness or abundance overall, 13 of 18 guilds (richness and abundance) and 16 of 22 species analyzed. Five of 10 farming practice variables (including herbicide use, organic farm type) and 9 of 13 habitat variables (including hedgerow length, proportion of hay) were significant in best models. Risk modeling indicated that herbicide use could decrease primary grassland birds by one species (35% decline from 3.4 to 2.3 species) per site. Organic farming could benefit species of conservation concern by 49% (an increase from 7.6 to 11.4 grassland birds). An addition of 63 m of hedgerow could increase abundance and richness of short distance migrants by 50% (3.0 to 4.8 and 1.3 to 2.0, respectively). Increasing the proportion of hay on nonorganic farms to 50% could increase abundance of primary grassland bird by 40% (6.7 to 9.4). Our results provide support for alternative farmland designs and agricultural management systems that could enhance select bird species in farmland.

A Demographic Model to Evaluate Population Declines in the Endangered Streaked Horned Lark

The Streaked Horned Lark (Eremophila alpestris strigata) is listed as endangered by the State of Washington, USA and by Canada under the Species at Risk Act and is also classified as a federal candidate for listing under the Endangered Species Act in the USA. A substantial portion of Streaked Horned Lark habitat has been lost or degraded, and range contraction has occurred in Oregon, Washington, and British Columbia. We estimate the vital rates (fecundity, adult and juvenile survival) and population growth rate (λ) for Streaked Horned Larks breeding in Washington, USA and conduct a Life-Stage Simulation Analysis (LSA) to evaluate which vital rate has the greatest influence on λ. We simulated changes in the three vital rates to examine how much they would need to be adjusted either independently or in concert to achieve a stable Streaked Horned Lark population (λ = 1). We also evaluated which fecundity component (the number of fledglings per egg laid or renesting interval) had the greatest impact on λ. The estimate of population growth suggests that Streaked Horned Larks in Washington are declining rapidly (λ = 0.62 ± 0.10) and that local breeding sites are not sustainable without immigration. The LSA results indicate that adult survival had the greatest influence on λ, followed by juvenile survival and fecundity. However, increases in vital rates led to λ = 1 only when adult survival was raised from 0.47 to 0.85, juvenile survival from 0.17 to 0.58, and fecundity from 0.91 to 3.09. Increases in breeding success and decreases in the renesting interval influenced λ similarly; however, λ did not reach 1 even when breeding success was raised to 100% or renesting intervals were reduced to 1 day. Only when all three vital rates were increased simultaneously did λ approach 1 without requiring highly unrealistic increases in each vital rate. We conclude that conservation activities need to target all or multiple vital rates to be successful. The baseline data presented here and subsequent efforts to manage Streaked Horned Larks will provide valuable information for management of other declining Horned Lark subspecies and other grassland songbirds across North America.

Patch Scale Nest-Site Selection by Marbled Murrelets (Brachyramphus marmoratus)

The Marbled Murrelet (Brachyramphus marmoratus) is a threatened alcid that nests almost exclusively in old-growth forests along the Pacific coast of North America. Nesting habitat has significant economic importance. Murrelet nests are extremely difficult and costly to find, which adds uncertainty to management and conservation planning. Models based on air photo interpretation of forest cover maps or assessments by low-level helicopter flights are currently used to rank presumed Marbled Murrelet nesting habitat quality in British Columbia. These rankings are assumed to correlate with nest usage and murrelet breeding productivity. Our goal was to find the models that best predict Marbled Murrelet nesting habitat in the ground-accessible portion of the two regions studied. We generated Resource Selection Functions (RSF) using logistic regression models of ground-based forest stand variables gathered at plots around 64 nests, located using radio-telemetry, versus 82 random habitat plots. The RSF scores are proportional to the probability of nests occurring in a forest patch. The best models differed somewhat between the two regions, but include both ground variables at the patch scale (0.2-2.0 ha), such as platform tree density, height and trunk diameter of canopy trees and canopy complexity, and landscape scale variables such as elevation, aspect, and slope. Collecting ground-based habitat selection data would not be cost-effective for widespread use in forestry management; air photo interpretation and low-level aerial surveys are much more efficient methods for ranking habitat suitability on a landscape scale. This study provides one method for ground-truthing the remote methods, an essential step made possible using the numerical RSF scores generated herein.

Costs of Reproduction in Breeding Female Mallards: Predation Risk during Incubation Drives Annual Mortality

The effort expended on reproduction may entail future costs, such as reduced survival or fecundity, and these costs can have an important influence on life-history optimization. For birds with precocial offspring, hypothesized costs of reproduction have typically emphasized nutritional and energetic investments in egg formation and incubation. We measured seasonal survival of 3856 radio-marked female Mallards (Anas platyrhynchos) from arrival on the breeding grounds through brood-rearing or cessation of breeding. There was a 2.5-fold direct increase in mortality risk associated with incubating nests in terrestrial habitats, whereas during brood-rearing when breeding females occupy aquatic habitats, mortality risk reached seasonal lows. Mortality risk also varied with calendar date and was highest during periods when large numbers of Mallards were nesting, suggesting that prey-switching behaviors by common predators may exacerbate risks to adults in all breeding stages. Although prior investments in egg laying and incubation affected mortality risk, most relationships were not consistent with the cost of reproduction hypothesis; birds with extensive prior investments in egg production or incubation typically survived better, suggesting that variation in individual quality drove both relationships. We conclude that for breeding female Mallards, the primary cost of reproduction is a fixed cost associated with placing oneself at risk to predators while incubating nests in terrestrial habitats.

Investigating Targets of Avian Habitat Management to Eliminate an Ecological Trap

Ecological traps are attractive population sinks created when anthropogenic habitat alteration inadvertently creates a mismatch between the attractiveness of a habitat based upon its settlement cues, and its current value for survival or reproduction. Traps represent a new threat to the conservation of native species, yet little attention has been given to developing practical approaches to eliminating them. In the northern Rocky Mountains of Montana, Olive-sided Flycatchers (Contopus cooperi) prefer to settle in patches of selectively harvested forest versus burned forest despite the lower reproductive success and higher nest predation risk associated with the former habitat. I investigated characteristics of preferred perch sites for this species and how these preferences varied between habitats and sexes. I then built on previous research to develop a range of management prescriptions for reducing the attractiveness of selectively harvested forest, thereby disarming the ecological trap. Female flycatchers preferred to forage from shorter perch trees than males, and females’ perches were shorter than other available perch trees. Both sexes preferred standing dead perch trees (snags) and these preferences were most obvious in harvested forest where snags are rarer. Because previous research shows that snag density is linked to habitat preference and spruce/fir trees are preferred nest substrate, my results suggest these two habitat components are focal habitat selection cues. I suggest alternative and complementary strategies for eliminating the ecological trap for Olive-sided Flycatchers including: (1) reduced retention and creation of snags, (2) avoiding selective harvest in spruce, fir, and larch stands, (3) avoiding retention of these tree species, and (4) selecting only even-aged canopy height trees for retention so as to reduce perch availability for female flycatchers. Because these strategies also have potential to negatively impact habitat suitability for other forest species or even create new ecological traps, we urge caution in the application of our management recommendations.