Potential Role of Spruce Budworm in Range-Wide Decline of Canada Warbler

ABSTRACT The Canada warbler (Wilsonia canadensis) is one of many common neotropical migrants whose populations are in decline across their range. Influences of habitat loss and degradation on breeding or wintering grounds have been postulated as possible causes, but few empirical data exist to support a specific cause. Based on previous studies linking abundances of Canada warbler and spruce budworm (Choristoneura fumiferana), we hypothesized that the Canada warbler may be influenced by a persistent decline in spruce budworm throughout the bird's breeding range, a hypothesis that has received little attention. This hypothesis makes 5 predictions: 1) budworm outbreaks and warbler detections should be spatially and temporally coincident; 2) the relationship between Canada warbler and spruce budworm outbreaks should be similar to relationships for other warblers known to be spruce budworm associates; 3) the relationship should be stronger than for warblers lacking an association with spruce budworm; 4) because temporal trends of both spruce budworm and Canada warblers have varied throughout Canadian provinces, declines in Canada warblers should be seen only in provinces where spruce budworm also declined; and 5) variation in Canada warbler abundance should reflect variation in supply of preferred habitat for the spruce budworm if habitat rather than budworm abundance is the key. Our analyses supported predictions 1–4, suggesting that Canada warbler may be even more closely associated with spruce budworm than are known associated species, a phenomenon noted in the literature but previously unexplained. Prediction 5 was not supported, because budworm habitat (area of mature and older balsam fir [Abies balsamea] and white spruce [Picea glauca]) remained constant in Ontario while warbler abundance declined. Although the correlative nature of these results precludes inference of a causal relationship between the declines of the Canada warbler and spruce budworm, we postulate that potential links may exist directly, where spruce budworm outbreaks provide elevated levels of insect prey items for breeding Canada warblers, or indirectly through changes in forest structure and composition following outbreaks. These results have implications when considering long-term trends in Canada warbler populations, because it may be impossible to alter population trends for species linked to the timing and magnitude of spruce budworm outbreaks.     

The interacting effects of food,spring temperature,and global climate cycles on population dynamics of a migratory songbird

Although long‐distance migratory songbirds are widely believed to be at risk from warming temperature trends, species capable of attempting more than one brood in a breeding season could benefit from extended breeding seasons in warmer springs. To evaluate local and global factors affecting population dynamics of the black‐throated blue warbler (Setophaga caerulescens), a double‐brooded long‐distance migrant, we used Pradel models to analyze 25 years of mark–recapture data collected in New Hampshire, USA. We assessed the effects of spring temperature (local weather) and the El Niño Southern Oscillation index (a global climate cycle), as well as predator abundance, insect biomass, and local conspecific density on population growth in the subsequent year. Local and global climatic conditions affected warbler populations in different ways. We found that warbler population growth was lower following El Niño years (which have been linked to poor survival in the wintering grounds and low fledging weights in the breeding grounds) than La Niña years. At a local scale, populations increased following years with warm springs and abundant late‐season food, but were unaffected by spring temperature following years when food was scarce. These results indicate that the warming temperature trends might have a positive effect on recruitment and population growth of black‐throated blue warblers if food abundance is sustained in breeding areas. In contrast, potential intensification of future El Niño events could negatively impact vital rates and populations of this species.     

Landscape host abundance and configuration regulate periodic outbreak behavior in spruce budworm Choristoneura fumiferana

Landscape‐level forest management has long been hypothesized to affect forest insect outbreak dynamics, but empirical evidence remains elusive. We hypothesized that the combination of increased hardwood relative to host tree species, prevalence of younger forests, and fragmentation of those forests due to forest harvesting legacies would reduce outbreak intensity, increase outbreak frequency, and decrease spatial synchrony in spruce budworm Choristoneura fumiferana outbreaks. We investigated these hypotheses using tree ring samples collected across 51 sites pooled into 16 subareas distributed across a large ecoregion spanning the international border between Ontario (Canada), and Minnesota (USA). This ecoregion contains contrasting land management zones with clear differences in forest landscape structure (i.e. forest composition and spatial configuration) while minimizing the confounding influence of climate. Cluster analyses of the 76‐yr time‐series generally grouped by subareas found within the same land management zone. Spatial nonparametric covariance analysis indicated that the highest and lowest degree of spatial synchrony of spruce budworm outbreaks were found within unmanaged wilderness and lands managed at fine spatial scales in Minnesota, respectively. Using multivariate analysis, we also found that forest composition, configuration, and climate together accounted for a total of 40% of the variance in outbreak chronologies, with a high level of shared variance between composition and configuration (13%) and between composition and climate (9%). At the scale of our study, climate on its own did not explain any of the spatial variation in outbreaks. Outbreaks were of higher frequency, lower intensity, and less spatially synchronized in more fragmented, younger forests with a lower proportion of host species, with opposing outbreak characteristics observed in regions characterised by older forests with more concentrated host species. Our study is the first quantitative evaluation of the long‐standing ‘silvicultural hypothesis’ of spruce budworm management specifically conducted at a spatio‐temporal scale for which it was intended.     

Breeding timed to maximize reproductive success for a migratory songbird: the importance of phenological asynchrony

Phenological advances and trophic mismatches are frequently reported ecological consequences of climate warming. Trophic mismatches occur when phenological responses to environmental conditions differ among trophic levels such that the timing of resource demand by consumers becomes decoupled from supply. We used 25 years of demographic measurements of a migratory songbird (the black‐throated blue warbler Setophaga caerulescens) to compare its breeding phenology to the phenology of both its caterpillar prey and the foliage on which caterpillars feed. Caterpillar biomass in this forest did not show a predictable seasonal pulse. Nest initiation by warblers in this northern hardwood forest was therefore not timed to coincide with a peak in food availability for nestlings. Nonetheless, timing of first clutches was strongly associated with spring leaf expansion (slope ± SE = 0.56 ± 0.08 days per day of change in leaf phenology, R² = 0.66). Warblers adjusted the timing of breeding to early springs mainly by shortening the interval between arrival and clutch initiation, but this likely has limits because recent early springs are approaching the relatively inflexible dates when birds arrive on the breeding grounds. Although the timing of first nests did not match 1:1 with leaf‐out phenology, the adjustments in breeding time maximized mean annual reproductive success. Nest predation had the greatest effect on annual reproductive success, but the ability of nesting warblers to appropriately track leaf phenology accounted for effects on annual reproductive success comparable to the influence of variation in caterpillar abundance and conspecific density. Nesting phenology in black‐throated blue warblers was generally well matched to the timing of leaf‐out, even though the match was not 1:1. Without measurements of reproductive success, these unequal phenological shifts might otherwise have been interpreted as having negative ecological consequences.     

Width of riparian buffer and structure of adjacent plantations influence occupancy of conservation priority birds

Conservation of biodiversity on forest landscapes dominated by plantations has become an increasingly important topic, and opportunities to maintain or enhance biodiversity within these forests need to be recognized and applied. Riparian buffers of mature forest retained along streams in managed forest landscapes offer an opportunity to enhance biodiversity across these landscapes. However, influence of buffer width and structure of adjacent plantations on habitat use by birds is not well understood. We modeled probability of occupancy, while accounting for variable detection probabilities, for 16 bird species of regional conservation importance in the Ouachita Mountains of Arkansas, USA. We examined occurrence of breeding birds in streamside management zone (SMZ) buffers embedded in three structural classes of pine plantation: young open-canopy, closed-canopy, and older thinned plantations. Our occupancy models included a positive association with SMZ width for nine bird species associated with mature forests. Models for three early successional species (prairie warbler [Dendroica discolor], white-eyed vireo [Vireo griseus], and northern bobwhite [Colinus virginianus]) included a negative association with SMZ width. Occupancy models for Acadian flycatcher (Empidonax virescens), summer tanager (Piranga rubra), pine warbler (Dendroica pinus), prairie warbler, and northern bobwhite also included structural condition of adjacent plantations, but most species did not appear affected by condition of surrounding plantations. We found diverse responses among species to width of retained SMZs and structure of adjacent plantations; some species apparently benefitted from SMZs >100 m wide, while others benefitted from narrow buffers. Furthermore, most species traditionally associated with mature forests were common in narrow SMZs, regardless of width. Thus, optimal width of SMZs relative to avian conservation depends on the species of greatest conservation interest.     

Influence of forest structure on density and nest success of mature forest birds in managed landscapes

Managing for forest wildlife requires attention not only to quantity but quality of forests within the landscape. We examined the extent to which local structural attributes and landscape context of forest stands explained variation in density and reproductive success of mature forest birds across 12 sites in southeast Ohio, USA, 2004–2006. Results suggest that several structural characteristics influenced bird–habitat relationships in our study. Densities of 3 songbird species (i.e., ovenbird [Seiurus aurocapilla], cerulean warbler [Setophaga cerulea], and scarlet tanager [Piranga olivacea]) were positively related to canopy openness, which is usually a function of canopy gaps. Habitat attributes described by ground litter, understory density, and canopy height were positively associated with densities of ground (i.e., worm-eating warbler [Helmitheros vermivorum]), or shrub nesting species (i.e., Kentucky and hooded warblers [Geothlypis formosa and Setophaga citrina], respectively). Furthermore, the number of small trees likely drove the positive relationship between density of wood thrush (Hylocichla mustelina), a subcanopy nester. After accounting for temporal variability in daily nest survival rates, the odds of nest survival for all species increased 10.5% for every 1% increase in canopy openness and decreased 1.4% for each 5% increase in understory vegetation density. Habitat–nest survival relationships were not apparent at the level of the individual species. Our results suggest that structural attributes produced by increasing habitat heterogeneity may be necessary for conservation of forest bird communities. © 2012 The Wildlife Society.     

Reservoir water levels do not influence daily mass gain of warblers at a riparian stopover site

ABSTRACT Hydroelectric dam operations that lead to fluctuations in the water levels of reservoirs can influence the amount of riparian habitat available for migrating songbirds and may impact the use and quality of remaining habitat. Our objective was to determine if use of riparian habitats and mass gain by five warbler species at the Columbia River‐Revelstoke Migration Monitoring Station in British Columbia, Canada, were influenced by water levels in the surrounding Arrow Lakes Reservoir. We analyzed fall migration data collected from 1998 to 2006. Capture rates of American Redstarts (Setophaga ruticilla), Common Yellowthroats (Geothlypis trichas), Orange‐crowned Warblers (Vermivora celata), Wilson's Warblers (Wilsonia pusilla), and Yellow Warblers (Dendroica petechia) varied between years and weeks of the migration period, but were not affected by annual or weekly variation in water levels. Annual variation in capture rates was driven by hatch‐year birds (>80% of individuals captured were juveniles) and could reflect conditions on the breeding grounds that influence productivity. We found that mass gain by the five species of warblers varied between 0.32% and 0.98% of lean body mass/hour. Mass gain did not vary between years or across weeks of the migration period and was not influenced by annual or weekly variation in reservoir water levels. Although the amount of available riparian habitat was reduced when reservoir water levels were high, we found no evidence that this loss of habitat influenced either the number of warblers or the mass gain of warblers using the riparian habitat that remained. Body mass at the time of first capture varied between years and across weeks for all five species. For American Redstarts and Orange‐crowned Warblers, body mass declined as average weekly water levels increased, a pattern that could arise if water levels influenced either their settlement decisions or length of stay.     

Detection range of songbirds using a stopover site by automated radio‐telemetry

A major uncertainty in automated radio‐telemetry studies of small birds is the detection range of receiving antennas. We compared simultaneous daytime detections (± 30 s) by automated and manual radio‐telemetry to assess detection probability and the proportion of transmissions detected for birds on migratory stopover as a function of distance, foraging guild (Black‐throated Blue Warblers, Setophaga caerulescens, and Yellow‐rumped Warblers, Dendroica coronata coronata, represented mid‐canopy foliage gleaners and White‐throated Sparrows, Zonotrichia albicollis, represented a ground forager), habitat type, meteorological variables, tower antenna number (1–4), and the position of a bird relative to the receiving antenna's bearing (offset angle). Our study was conducted at a migratory stopover site in southern Ontario, Canada. Most detections were in dense to sparse forest, and all individuals were within 1.03 km of the automated receiving station. Daily detection probability was near 100% for both foraging guilds. However, within 30 s before and after a manual radio‐telemetry location was made, detection probability and the proportion of transmissions detected by automated radio‐telemetry declined with distance, was higher for warblers than sparrows, and was lowest for 90° offset angles. Our results suggest that when research goals do not require detections with high temporal frequency, e.g., estimation of departure date or daily departure probability, our study design had an effective detection range of at least 1 km. However, where temporal precision is required, e.g., to investigate movements and changes in activity levels during stopover, detection range was ~300 m for ground‐foraging sparrows and 600 m for mid‐canopy foraging warblers, which is much lower than the presumed detection range of antennas under optimal conditions (15 km). This corresponds to a spatial area of coverage for forest‐dwelling birds of ~0.3–1.1 km². Our results suggest that to optimally configure an automated radio‐telemetry array at the regional scale, investigators should carefully consider detection range and its underlying covariates, including species type, the habitat matrix, and the orientation of antennas relative to preferred habitat.     

A dendroecological reconstruction of western spruce budworm outbreaks (Choristoneura occidentalis) in the Front Range,Colorado, from 1720 to 1986

Radial increment cores from Douglas-fir (Pseudotsuga menziesii) and blue spruce (Picea pungens), defoliated by western spruce budworm (Choristoneura occidentalis), were analyzed by means of dendrochronological methods and compared with cores from undefoliated ponderosa pine (Pinus ponderosa) and lodgepole pine (Pinus contorta) growing on the same sites in the Front Range, Colorado. Extensive deforestation during the gold and silver booms in the second part of the nineteenth century led to dense and almost pure stands of shadetolerant budworm host species. By using the skeleton plot method, the number of trees with clear growth reductions is obtained, thus representing an exact record of forest insect attacks. The analysis of abrupt growth reductions revealed at least nine outbreaks of western spruce budworm between 1720 and 1986, the majority occurring in the nineteenth century. The outbreaks were graphically compared with periods of attack in New Mexico and Colorado which were detected by other scientists employing tree-ring measurement techniques. No increase in the frequency of severe outbreaks during the twentieth century was observed, yet there is some evidence that the most recent outbreak might be the most severe ever recorded. Open Douglas-fir stands on higher sites were more susceptible to heavy budworm attack than dense stands on lower sites. Blue spruce was less frequently and less severely attacked than Douglas-fir. The spatial pattern of historical outbreaks generally was very patchy.     

Early singers attend to conspecific but not heterospecific behavioural cues at dawn

During the dawn chorus songbirds initiate singing activities just prior to sunrise. Environmental factors affect the timing of the dawn chorus, but relatively little is known about how behavioural cues influence chorus timing. We assessed whether early playback of conspecific and heterospecific song influenced chorus start times in two early‐singing temperate species, hermit thrush Catharus guttatus and veery C. fuscescens. We used arrays of GPS time‐synchronized recorders to record the dawn chorus at 30 locations. In each location, thirty minutes prior to natural chorus initiation, we broadcast: 1) an early singing species (hermit thrush or veery), 2) a late singing species (black‐throated green warbler Setophaga virens or Nashville warbler Oreothlypis ruficapilla), 3) a noise control, and 4) also recorded without playback (silent control). Playback treatments were separated by two‐three days at each location. Both hermit thrush and veery sang significantly earlier in response to conspecific playback compared to noise or silent controls. Both species also sang earlier in response to conspecific playback compared to the day before or the day after playback. Neither species sang earlier in response to heterospecific playback (regardless of whether the broadcast species was a naturally early or late song initiator) compared to noise or silent controls. Our results support the theory that the dawn chorus is a communication network where individuals attend primarily to conspecific cues; heterospecific song appears to have minimal impacts on the chorus start times of the two early‐singing species we investigated.     

Gap‐crossing decisions of forest birds in a fragmented landscape

Habitat loss and fragmentation are recognized as primary drivers of biodiversity loss worldwide. To understand the functional effects of habitat fragmentation on bird populations, data on movement across gaps in habitat cover are necessary, although rarely available. In this study, we used call playback to simulate a conspecific territorial intruder to entice birds to move through the landscape in a predictable and directional manner. We then quantified the probability of movement in continuous forest and across cleared gaps for two forest‐dependent species, the grey shrike‐thrush (Colluricincla harmonica) and the white‐throated treecreeper (Cormobates leucophaeus). Fifty‐four playback trials were conducted for each species across distances ranging from 25 to 480 m in continuous forest and 15–260 m across gaps in a forest‐agricultural landscape in southern Victoria, Australia. The probability of movement was significantly reduced by gaps in forest cover for both species. Shrike‐thrushes were six times more likely to move 170 m in continuous forest than to cross 170‐m gaps. The mean probability that treecreepers would cross any gap at all was less than 0.5, and they were three times less likely to move 50 m across a gap than through continuous forest. Both species displayed non‐linear responses to increasing gap distance: we identified a gap‐tolerance threshold of 85 m for the shrike‐thrush and 65 m for the treecreeper beyond which individuals were most unlikely to cross. The presence of scattered paddock trees increased functional connectivity for the shrike‐thrush, with individuals crossing up to 260 m when scattered trees were present. We conclude that gaps in habitat cover are barriers to movement, and that characteristics of the intervening matrix influence landscape permeability.     

Continent‐wide population genomic structure and phylogeography of North America’s most destructive conifer defoliator,the spruce budworm (Choristoneura fumiferana)

The spruce budworm, Choristoneura fumiferana, is presumed to be panmictic across vast regions of North America. We examined the extent of panmixia by genotyping 3,650 single nucleotide polymorphism (SNP) loci in 1975 individuals from 128 collections across the continent. We found three spatially structured subpopulations: Western (Alaska, Yukon), Central (southeastern Yukon to the Manitoba–Ontario border), and Eastern (Manitoba–Ontario border to the Atlantic). Additionally, the most diagnostic genetic differentiation between the Central and Eastern subpopulations was chromosomally restricted to a single block of SNPs that may constitute an island of differentiation within the species. Geographic differentiation in the spruce budworm parallels that of its principal larval host, white spruce (Picea glauca), providing evidence that spruce budworm and spruce trees survived in the Beringian refugium through the Last Glacial Maximum and that at least two isolated spruce budworm populations diverged with spruce/fir south of the ice sheets. Gene flow in the spruce budworm may also be affected by mountains in western North America, habitat isolation in West Virginia, regional adaptations, factors related to dispersal, and proximity of other species in the spruce budworm species complex. The central and eastern geographic regions contain individuals that assign to Eastern and Central subpopulations, respectively, indicating that these barriers are not complete. Our discovery of previously undetected geographic and genomic structure in the spruce budworm suggests that further population modelling of this ecologically important insect should consider regional differentiation, potentially co‐adapted blocks of genes, and gene flow between subpopulations.     

Long‐term successional forest dynamics: species and community responses to climatic variability

Question: Are trees sensitive to climatic variability, and do tree species differ in their responses to climatic variability? Does sensitivity of forest communities to climatic variability depend on stand composition? Location: Mixed young forest at Walker Branch Watershed near Oak Ridge, East Tennessee, USA. Methods: Using a long‐term dataset (1967–2006), we analyzed temporal forest dynamics at the tree and species level, and community dynamics for forest stands that differed in initial species composition (i.e., chestnut oak, oak–hickory, pine, and yellow poplar stands). Using summer drought and growing season temperature as defined climate drivers, we evaluated relationships between forest dynamics and climate across levels of organization. Results: Over the four‐decade study period, forest communities underwent successional change and substantially increased in biomass. Variation in summer drought and growing season temperature contributed to temporal biomass dynamics for some tree species, but not for others. Stand‐level responses to climatic variability were related to the responses of component species, except in pine stands. Pinus echinata, the dominant species in pine stands, decreased over time due to periodic outbreaks of pine bark beetle (Dendroctonus frontalis). These outbreaks at Walker Branch could not be directly related to climatic conditions. Conclusions: The results indicate that sensitivity of developing forests to climatic variability is stand type‐dependent, and hence is a function of species composition. However, in the long term, direct effects of climatic variability on forest dynamics may be small relative to autogenic successional processes or climate‐related insect outbreaks. Empirical studies testing for interactions between forest succession and climatic variability are needed.     

Polymorphic microsatellite loci from the American goldfinch (Carduelis tristis) and their cross‐amplification in a variety of passerine species

Cardueline finches have become important models in studies of sexual selection. Here I describe eight polymorphic microsatellite loci isolated from the American goldfinch (Carduelis tristis) that are useful in parentage and heterozygosity studies. The number of alleles per locus ranged from eight to 38 based on 252 goldfinches. The markers are polymorphic in at least four other Cardueline finch species, and several amplify in nonfinch passerines including red‐winged blackbird (Agelaius phoeniceus), black‐throated blue warbler (Dendroica caerulescens) and tufted titmouse (Baeolophus bicolor), indicating they should be useful in paternity and population studies of a variety of species.     

Forest defoliator outbreaks under climate change: effects on the frequency and severity of outbreaks of five pine insect pests

To identify general patterns in the effects of climate change on the outbreak dynamics of forest‐defoliating insect species, we examined a 212‐year record (1800–2011) of outbreaks of five pine‐defoliating species (Bupalus piniarius, Panolis flammea, Lymantria monacha, Dendrolimus pini, and Diprion pini) in Bavaria, Germany for the evidence of climate‐driven changes in the severity, cyclicity, and frequency of outbreaks. We also accounted for historical changes in forestry practices and examined effects of past insecticide use to suppress outbreaks. Analysis of relationships between severity or occurrence of outbreaks and detrended measures of temperature and precipitation revealed a mixture of positive and negative relationships between temperature and outbreak activity. Two moth species (P. flammea and Dendrolimus pini) exhibited lower outbreak activity following years or decades of unusually warm temperatures, whereas a sawfly (Diprion pini), for which voltinism is influenced by temperature, displayed increased outbreak occurrence in years of high summer temperatures. We detected only one apparent effect of precipitation, which showed Dendrolimus pini outbreaks tending to follow drought. Wavelet analysis of outbreak time series suggested climate change may be associated with collapse of L. monacha and Dendrolimus pini outbreak cycles (loss of cyclicity and discontinuation of outbreaks, respectively), but high‐frequency cycles for B. piniarius and P. flammea in the late 1900s. Regional outbreak severity was generally not related to past suppression efforts (area treated with insecticides). Recent shifts in forestry practices affecting tree species composition roughly coincided with high‐frequency outbreak cycles in B. piniarius and P. flammea but are unlikely to explain the detected relationships between climate and outbreak severity or collapses of outbreak cycles. Our results highlight both individualistic responses of different pine‐defoliating species to climate changes and some patterns that are consistent across defoliator species in this and other forest systems, including collapsing of population cycles.     

Insect outbreaks increase populations and facilitate reproduction in a cavity‐dependent songbird,the Mountain Chickadee Poecile gambeli

Large‐scale insect outbreaks, typical of temperate ecosystems, may increase food availability and influence species interactions in insectivorous bird communities. We assessed how population densities of a secondary cavity nester, the Mountain Chickadee Poecile gambeli, varied with densities of avian cavity excavators, potential competitors and nest predators, during two large‐scale insect outbreaks of Western Spruce Budworm Choristoneura occidentalis and Mountain Pine Beetle Dendroctonus ponderosae. At the regional level, Mountain Chickadee densities doubled with increases in budworm and beetle availability, then declined with decreases in beetle availability, despite high budworm supply. At the site level, densities of excavators were the best predictor of site‐level variation in Chickadee densities in the following year. During and after the beetle outbreak, Mountain Chickadees used more cavities excavated by Red‐breasted Nuthatches Sitta canadensis and Downy Woodpeckers Picoides pubescens, both of which are bark insectivores that increased in densities concomitantly with the beetle outbreak and whose foraging activities may have facilitated the Mountain Chickadees' access to bark beetles. Thus, Mountain Chickadees showed a numerical response to the food pulse at the regional level, but a functional response to the pulse of nesting cavities at the site level. Plasticity in resource selection and heterospecific attraction may allow Mountain Chickadees to respond to resource pulses in highly variable environments.     

No short‐term change in avian assemblage following removal of Yellow‐throated Miner (Manorina flavigula) colonies

The overabundance of Yellow‐throated Miner (Manorina flavigula) has been shown to negatively affect the abundance and richness of small birds in areas they occupy, leading to homogenization of the avifauna across the fragmented landscape. In this study, we took advantage of a planned management cull to ask the question, does the removal of Yellow‐throated Miner colonies cause an immediate change in avian species richness and abundance? This cull was undertaken around the Bronzewing Flora and Fauna Reserve (north‐western Victoria, Australia) in order to protect a resident population of endangered Black‐eared Miner (M. melanotis) from hybridization. We conducted avian surveys along roadsides surrounding the reserve at Yellow‐throated Miner colonies (n = 6), control sites with no miners (n = 7), and where colonies were removed (n = 3). We found that the cull was followed by only a very modest increase in the species richness and abundance of small birds, with no significant effects on avian assemblage overall. This result contrasts with far more dramatic increases following culls of other species of miner. Sites where miners were removed were not depauperate of other species prior to the cull, which could have been due to a combination of proximity to refuge for small birds in a neighbouring reserve or the low numbers of miners that made up each culled colony. This study highlights that assumed effects of a management action may be highly dependent upon spatial and temporal context.     

Who is in the neighborhood? Conspecific and heterospecific responses to perceived density for breeding habitat selection

Theoretical models of habitat selection often incorporate negative density dependence. Despite strong negative density‐dependent effects on habitat selection, more recent studies indicate that animals settle near members of their own (conspecific) and other species (heterospecific) when selecting habitat with social cues. Social cue use for habitat selection is particularly common among songbirds, but few studies have investigated if songbirds use social cues to assess conspecific or heterospecific density (as opposed to just presence/absence) when making settlement decisions. We conducted a playback experiment to evaluate if yellow warblers (Setophaga petechia) and willow flycatchers (Empidonax traillii), two potential competitors for breeding habitat, use social cues to assess density (conspecific for warblers and heterospecific for flycatchers) when selecting breeding locations at two spatial scales. We simulated yellow warbler density to be high or low at multiple treatment plots (3.14 ha) with song playback and then evaluated settlement decisions by comparing yellow warbler and willow flycatcher abundances across plots (broad‐scale habitat selection) and individual space use within plots (fine‐scale territory establishment). Yellow warbler density treatments did not affect habitat selection by yellow warblers at the broad scale, but caused individuals to cluster territories at high‐density treatments. Willow flycatchers were most abundant at high‐density treatment plots, but yellow warbler density treatments did not affect territory locations. The results indicate that perceived density affects the habitat selection process for both conspecifics and heterospecifics.     

Estimating the Effects of Habitat and Biological Interactions in an Avian Community

We used repeated sightings of individual birds encountered in community-level surveys to investigate the relative roles of habitat and biological interactions in determining the distribution and abundance of each species. To analyze these data, we developed a multispecies N-mixture model that allowed estimation of both positive and negative correlations between abundances of different species while also estimating the effects of habitat and the effects of errors in detection of each species. Using a combination of single- and multispecies N-mixture modeling, we examined for each species whether our measures of habitat were sufficient to account for the variation in encounter histories of individual birds or whether other habitat variables or interactions with other species needed to be considered. In the community that we studied, habitat appeared to be more influential than biological interactions in determining the distribution and abundance of most avian species. Our results lend support to the hypothesis that abundances of forest specialists are negatively affected by forest fragmentation. Our results also suggest that many species were associated with particular types of vegetation as measured by structural attributes of the forests. The abundances of 6 of the 73 species observed in our study were strongly correlated. These species included large birds (American Crow (Corvus brachyrhynchos) and Red-winged Blackbird (Agelaius phoeniceus)) that forage on the ground in open habitats and small birds (Red-eyed Vireo (Vireo olivaceus), House Wren (Troglodytes aedon), Hooded Warbler (Setophaga citrina), and Prairie Warbler (Setophaga discolor)) that are associated with dense shrub cover. Species abundances were positively correlated within each size group and negatively correlated between groups. Except for the American Crow, which preys on eggs and nestlings of small song birds, none of the other 5 species is known to display direct interactions, so we suspect that the correlations may have been associated with species-specific responses to habitat components not adequately measured by our covariates.     

Experimental climate warming alters aspen and birch phytochemistry and performance traits for an outbreak insect herbivore

Climate change and insect outbreaks are key factors contributing to regional and global patterns of increased tree mortality. While links between these environmental stressors have been established, our understanding of the mechanisms by which elevated temperature may affect tree–insect interactions is limited. Using a forest warming mesocosm, we investigated the influence of elevated temperature on phytochemistry, tree resistance traits, and insect performance. Specifically, we examined warming effects on forest tent caterpillar (Malacosoma disstria) and host trees aspen (Populus tremuloides) and birch (Betula papyrifera). Trees were grown under one of three temperature treatments (ambient, +1.7 °C, +3.4 °C) in a multiyear open‐air warming experiment. In the third and fourth years of warming (2011, 2012), we assessed foliar nutrients and defense chemistry. Elevated temperatures altered foliar nitrogen, carbohydrates, lignin, and condensed tannins, with differences in responses between species and years. In 2012, we performed bioassays using a common environment approach to evaluate plant‐mediated indirect warming effects on larval performance. Warming resulted in decreased food conversion efficiency and increased consumption, ultimately with minimal effect on larval development and biomass. These changes suggest that insects exhibited compensatory feeding due to reduced host quality. Within the context of observed phytochemical variation, primary metabolites were stronger predictors of insect performance than secondary metabolites. Between‐year differences in phytochemical shifts corresponded with substantially different weather conditions during these two years. By sampling across years within an ecologically realistic and environmentally open setting, our study demonstrates that plant and insect responses to warming can be temporally variable and context dependent. Results indicate that elevated temperatures can alter phytochemistry, tree resistance traits, and herbivore feeding, but that annual weather variability may modulate warming effects leading to uncertain consequences for plant–insect interactions with projected climate change.