Simulated treatment effects on bird communities inform landscape-scale dry conifer forest management.

Human land use and climate change have increased forest density and wildfire risk in dry conifer forests of western North America, threatening various ecosystem services, including habitat for wildlife. Government policy supports active management to restore historical structure and ecological function. Information on potential contributions of restoration to wildlife habitat can allow assessment of tradeoffs with other ecological benefits when prioritizing treatments. We predicted avian responses to simulated treatments representing alternative scenarios to inform landscape-scale forest management planning along the Colorado Front Range. We used data from the Integrated Monitoring in Bird Conservation Regions program to inform a hierarchical multispecies occupancy model relating species occupancy and richness with canopy cover at two spatial scales. We then simulated changes in canopy cover (remotely sensed in 2018) under three alternative scenarios, (1) a "fuels reduction" scenario representing landscape-wide 30% reduction in canopy cover, (2) a "restoration" scenario representing more nuanced, spatially variable treatments targeting historical conditions, and (3) a reference, no-change scenario. Model predictions showed areas of potential gains and losses for species richness, richness of ponderosa pine forest habitat specialists, and the ratio of specialists to generalists at two (1 km2 and 250 m2 ) spatial scales. Under both fuels reduction and restoration scenarios, we projected greater gains than losses for species richness. Surprisingly, despite restoration more explicitly targeting ecologically relevant historical conditions, fuels reduction benefited bird species richness over a greater spatial extent than restoration, particularly in the lower montane life zone. These benefits reflected generally positive species associations with moderate canopy cover promoted more consistently under the fuels reduction scenario. In practice, contemporary forest management is likely to lie somewhere between the fuels reduction and restoration scenarios represented here. Therefore, our results inform where and how active forest management can best support avian diversity. Although our study raises questions regarding the value of including landscape-scale heterogeneity as a management objective, we do not question the value of targeting finer scale heterogeneity (i.e., stand and treatment level). Rather, our results combined with those from previous work clarify the scale at which targeting structural heterogeneity and historical reference conditions can promote particular ecosystem services.

Development and evaluation of habitat suitability models for nesting white-headed woodpecker (Dryobates albolarvatus) in burned forest.

Salvage logging in burned forests can negatively affect habitat for white-headed woodpeckers (Dryobates albolarvatus), a species of conservation concern, but also meets socioeconomic demands for timber and human safety. Habitat suitability index (HSI) models can inform forest management activities to help meet habitat conservation objectives. Informing post-fire forest management, however, involves model application at new locations as wildfires occur, requiring evaluation of predictive performance across locations. We developed HSI models for white-headed woodpeckers using nest sites from two burned-forest locations in Oregon, the Toolbox (2002) and Canyon Creek (2015) fires. We measured predictive performance by developing one model at each of the two locations and quantifying discrimination of nest from reference sites at two other wildfire locations where the model had not been developed (either Toolbox or Canyon Creek, and the Barry Point Fire [2011]). We developed and evaluated Maxent models based on remotely sensed environmental metrics to support habitat mapping, and weighted logistic regression (WLR) models that combined remotely sensed and field-collected metrics to inform management prescriptions. Both Maxent and WLR models developed either at Canyon Creek or Toolbox performed adequately to inform management when applied at the alternate Toolbox or Canyon Creek location, respectively (area under the receiver-operating-characteristic curve [AUC] range = 0.61-0.72) but poorly when applied at Barry Point (AUC = 0.53-0.57). The final HSI models fitted to Toolbox and Canyon Creek data quantified suitable nesting habitat as severely burned or open sites adjacent to lower severity and closed canopy sites, where foraging presumably occurs. We suggest these models are applicable at locations similar to development locations but not at locations resembling Barry Point, which were characterized by more (pre-fire) canopy openings, larger diameter trees, less ponderosa pine (Pinus ponderosa), and more juniper (Juniperus occidentalis). Considering our results, we recommend caution when applying HSI models developed at individual wildfire locations to inform post-fire management at new locations without first evaluating predictive performance.

Dry conifer forest restoration benefits Colorado Front Range avian communities.

Fire suppression has increased stand density and risk of severe, stand-replacing wildfire in lower elevation dry conifer forests of western North America, threatening ecological function. The U.S. Forest Service's Collaborative Forest Landscape Restoration Program (CFLRP) aims to mitigate impacts to ecological function, while mandating effectiveness monitoring to verify restoration success. Expected benefits include improved conditions for biodiversity, but relatively few empirical studies evaluate restoration effects on biodiversity. We applied the Integrated Monitoring in Bird Conservation Regions program to survey birds in relation to CFLRP treatments along the Colorado Front Range in 2015-2017. We employed hierarchical models to analyze species occupancy and richness at 1972 points nested within 141 1-km2 grid cells. Our objectives were to investigate (1) species occupancy relationships with treatments at local (point) and landscape (grid) spatial scales, (2) potential mechanisms for treatment relationships considering species and treatment relationships with forest structure and composition (i.e., habitat relationships), and (3) treatment and habitat relationships with species richness. The data supported positive and negative point-level treatment relationships, suggesting uneven species distributions between treated and untreated points. At the grid scale, however, we only found positive species relationships with percent area treated, and accordingly, grid-level species richness increased with treatment extent. Potential mechanisms for treatment relationships included treatments generating foraging opportunities for aerial insectivores by opening the canopy, improving conditions for ground-associated species by increasing herbaceous growth, and limiting opportunities for shrub-nesting species by reducing shrub cover. Landscape-scale patterns suggest CFLRP treatments can benefit avian communities by generating habitat for open-forest species without necessarily eliminating habitat for closed-forest species. Our results provide evidence for a commonly expected but rarely verified pattern of increased species richness with forest heterogeneity. We suggest restoration treatments will most benefit forest bird diversity by reducing canopy cover, encouraging herbaceous ground cover, limiting ladder fuel species, and encouraging shrub diversity in canopy openings, while maintaining some dense forest stands on the landscape.

Simulations inform design of regional occupancy-based monitoring for a sparsely distributed, territorial species.

Sparsely distributed species attract conservation concern, but insufficient information on population trends challenges conservation and funding prioritization. Occupancy-based monitoring is attractive for these species, but appropriate sampling design and inference depend on particulars of the study system. We employed spatially explicit simulations to identify minimum levels of sampling effort for a regional occupancy monitoring study design, using white-headed woodpeckers (Picoides albolvartus), a sparsely distributed, territorial species threatened by habitat decline and degradation, as a case study. We compared the original design with commonly proposed alternatives with varying targets of inference (i.e., species range, space use, or abundance) and spatial extent of sampling. Sampling effort needed to achieve adequate power to observe a long-term population trend (≥80% chance to observe a 2% yearly decline over 20 years) with the previously used study design consisted of annually monitoring ≥120 transects using a single-survey approach or ≥90 transects surveyed twice per year using a repeat-survey approach. Designs that shifted inference toward finer-resolution trends in abundance and extended the spatial extent of sampling by shortening transects, employing a single-survey approach to monitoring, and incorporating a panel design (33% of units surveyed per year) improved power and reduced error in estimating abundance trends. In contrast, efforts to monitor coarse-scale trends in species range or space use with repeat surveys provided extremely limited statistical power. Synthesis and applications. Sampling resolutions that approximate home range size, spatially extensive sampling, and designs that target inference of abundance trends rather than range dynamics are probably best suited and most feasible for broad-scale occupancy-based monitoring of sparsely distributed territorial animal species.

Ensemble modeling to predict habitat suitability for a large-scale disturbance specialist.

To conserve habitat for disturbance specialist species, ecologists must identify where individuals will likely settle in newly disturbed areas. Habitat suitability models can predict which sites at new disturbances will most likely attract specialists. Without validation data from newly disturbed areas, however, the best approach for maximizing predictive accuracy can be unclear (Northwestern U.S.A.). We predicted habitat suitability for nesting Black-backed Woodpeckers (Picoides arcticus; a burned-forest specialist) at 20 recently (≤6 years postwildfire) burned locations in Montana using models calibrated with data from three locations in Washington, Oregon, and Idaho. We developed 8 models using three techniques (weighted logistic regression, Maxent, and Mahalanobis D (2) models) and various combinations of four environmental variables describing burn severity, the north-south orientation of topographic slope, and prefire canopy cover. After translating model predictions into binary classifications (0 = low suitability to unsuitable, 1 = high to moderate suitability), we compiled "ensemble predictions," consisting of the number of models (0-8) predicting any given site as highly suitable. The suitability status for 40% of the area burned by eastside Montana wildfires was consistent across models and therefore robust to uncertainty in the relative accuracy of particular models and in alternative ecological hypotheses they described. Ensemble predictions exhibited two desirable properties: (1) a positive relationship with apparent rates of nest occurrence at calibration locations and (2) declining model agreement outside surveyed environments consistent with our reduced confidence in novel (i.e., "no-analogue") environments. Areas of disagreement among models suggested where future surveys could help validate and refine models for an improved understanding of Black-backed Woodpecker nesting habitat relationships. Ensemble predictions presented here can help guide managers attempting to balance salvage logging with habitat conservation in burned-forest landscapes where black-backed woodpecker nest location data are not immediately available. Ensemble modeling represents a promising tool for guiding conservation of large-scale disturbance specialists.

Effects of parents and Brown-headed Cowbirds (Molothrus ater) on nest predation risk for a songbird.

Nest predation limits avian fitness, so ornithologists study nest predation, but they often only document patterns of predation rates without substantively investigating underlying mechanisms. Parental behavior and predator ecology are two fundamental drivers of predation rates and patterns, but the role of parents is less certain, particularly for songbirds. Previous work reproduced microhabitat-predation patterns experienced by Yellow Warblers (Setophaga petechia) in the Mono Lake basin at experimental nests without parents, suggesting that these patterns were driven by predator ecology rather than predator interactions with parents. In this study, we further explored effects of post-initiation parental behavior (nest defense and attendance) on predation risk by comparing natural versus experimental patterns related to territory density, seasonal timing of nest initiation, and nest age. Rates of parasitism by Brown-headed Cowbirds (Molothrus ater) were high in this system (49% nests parasitized), so we also examined parasitism-predation relationships. Natural nest predation rates (NPR) correlated negatively with breeding territory density and nonlinearly (U-shaped relationship) with nest-initiation timing, but experimental nests recorded no such patterns. After adjusting natural-nest data to control for these differences from experimental nests other than the presence of parents (e.g., defining nest failure similarly and excluding nestling-period data), we obtained similar results. Thus, parents were necessary to produce observed patterns. Lower natural NPR compared with experimental NPR suggested that parents reduced predation rates via nest defense, so this parental behavior or its consequences were likely correlated with density or seasonal timing. In contrast, daily predation rates decreased with nest age for both nest types, indicating this pattern did not involve parents. Parasitized nests suffered higher rates of partial predation but lower rates of complete predation, suggesting direct predation by cowbirds. Explicit behavioral research on parents, predators (including cowbirds), and their interactions would further illuminate mechanisms underlying the density, seasonal, and nest age patterns we observed.

How avian nest site selection responds to predation risk: testing an 'adaptive peak hypothesis'.

1. Nest predation limits avian fitness, so birds should favour nest sites that minimize predation risk. Nevertheless, preferred nest microhabitat features are often uncorrelated with apparent variation in predation rates. 2. This lack of congruence between theory-based expectation and empirical data may arise when birds already occupy 'adaptive peaks'. If birds nest exclusively in low-predation microhabitats, microhabitat and nest predation may no longer be correlated even though predation ultimately shaped microhabitat selection. 3. This 'adaptive peak hypothesis' was tested for a population of Yellow Warblers (Dendroica petechia) focusing on two nest microhabitat features: concealment and height. Experimental nests measured relative predation risk both within and outside the microhabitat range typically occupied by natural nests to examine whether nest site choices made by birds restricted our ability to detect microhabitat effects on predation. 4. Within the natural range (30-80% concealment, >75 cm height), microhabitat-predation relationships were weak and inconsistent, and similar for experimental and natural nests. Over an extended range, however, experimental predation rates were elevated in exposed sites (<30% concealed), indicating a concealment-related 'adaptive plateau'. 5. Clay egg bite data revealed a concealment effect on avian predators, and the abundance of one avian predator group correlated with nest concealment among years, suggesting these predators may cue birds to modulate nest concealment choices. 6. This study demonstrates how avian responses to predation pressure can obscure the adaptive significance of nest site selection, so predation influences may be more important than apparent from published data.