Audio tagging of avian dawn chorus recordings in California, Oregon and Washington.

Background: Declines in biodiversity and ecosystem health due to climate change are raising urgent concerns. In response, large-scale multispecies monitoring programmes are being implemented that increasingly adopt sensor-based approaches such as acoustic recording. These approaches rely heavily on ecological data science. However, developing reliable algorithms for processing sensor-based data relies heavily on labelled datasets of sufficient quality and quantity. We present a dataset of 1,575 dawn chorus soundscape recordings, 141 being fully annotated (n = 32,994 annotations) with avian, mammalian and amphibian vocalisations. The remaining recordings were included to facilitate novel research applications. These recordings are paired with 48 site-level climatic, forest structure and topographic covariates. This dataset provides a valuable resource to researchers developing acoustic classification algorithms or studying biodiversity and wildlife behaviour and its relationship to environmental gradients. The dawn chorus recordings were collected as part of a long-term Northern Spotted Owl monitoring program; this demonstrates the complementary value of harnessing existing monitoring efforts to strengthen biodiversity sampling. New information: This dataset of dawn chorus soundscape recordings is one of the few open-access acoustic datasets annotated with non-biotic and both interspecific (across species) and intraspecific (within species) bird, mammal and amphibian sonotypes and the first that is paired with climatic, forest structure and topographical covariates extracted at recorder locations. This makes it a valuable resource for researchers studying the dawn chorus and its relationship to the environment.

Plant-hummingbird pollination networks exhibit limited rewiring after experimental removal of a locally abundant plant species.

Mutualistic relationships, such as those between plants and pollinators, may be vulnerable to the local extinctions predicted under global environmental change. However, network theory predicts that plant-pollinator networks can withstand species loss if pollinators switch to alternative floral resources (rewiring). Whether rewiring occurs following species loss in natural communities is poorly known because replicated species exclusions are difficult to implement at appropriate spatial scales. We experimentally removed a hummingbird-pollinated plant, Heliconia tortuosa, from within tropical forest fragments to investigate how hummingbirds respond to temporary loss of an abundant resource. Under the rewiring hypothesis, we expected that behavioural flexibility would allow hummingbirds to use alternative resources, leading to decreased ecological specialization and reorganization of the network structure (i.e. pairwise interactions). Alternatively, morphological or behavioural constraints-such as trait-matching or interspecific competition-might limit the extent to which hummingbirds alter their foraging behaviour. We employed a replicated Before-After-Control-Impact experimental design and quantified plant-hummingbird interactions using two parallel sampling methods: pollen collected from individual hummingbirds ('pollen networks', created from >300 pollen samples) and observations of hummingbirds visiting focal plants ('camera networks', created from >19,000 observation hours). To assess the extent of rewiring, we quantified ecological specialization at the individual, species and network levels and examined interaction turnover (i.e. gain/loss of pairwise interactions). H. tortuosa removal caused some reorganization of pairwise interactions but did not prompt large changes in specialization, despite the large magnitude of our manipulation (on average, >100 inflorescences removed in exclusion areas of >1 ha). Although some individual hummingbirds sampled through time showed modest increases in niche breadth following Heliconia removal (relative to birds that did not experience resource loss), these changes were not reflected in species- and network-level specialization metrics. Our results suggest that, at least over short time-scales, animals may not necessarily shift to alternative resources after losing an abundant food resource-even in species thought to be highly opportunistic foragers, such as hummingbirds. Given that rewiring contributes to theoretical predictions of network stability, future studies should investigate why pollinators might not expand their diets after a local resource extinction.

Climate-driven variation in dispersal ability predicts responses to forest fragmentation in birds.

Species sensitivity to forest fragmentation varies latitudinally, peaking in the tropics. A prominent explanation for this pattern is that historical landscape disturbance at higher latitudes has removed fragmentation-sensitive species or promoted the evolution of more resilient survivors. However, it is unclear whether this so-called extinction filter is the dominant driver of geographic variation in fragmentation sensitivity, particularly because climatic factors may also cause latitudinal gradients in dispersal ability, a key trait mediating sensitivity to habitat fragmentation. Here we combine field survey data with a morphological proxy for avian dispersal ability (hand-wing index) to assess responses to forest fragmentation in 1,034 bird species worldwide. We find that fragmentation sensitivity is strongly predicted by dispersal limitation and that other factors-latitude, body mass and historical disturbance events-have relatively limited explanatory power after accounting for species differences in dispersal. We also show that variation in dispersal ability is only weakly predicted by historical disturbance and more strongly associated with intra-annual temperature fluctuations (seasonality). Our results suggest that climatic factors play a dominant role in driving global variation in the impacts of forest fragmentation, emphasizing the need for more nuanced environmental policies that take into account local context and associated species traits.

Bee diversity decreases rapidly with time since harvest in intensively managed conifer forests.

Despite widespread concerns about the anthropogenic drivers of global pollinator declines, little information is available about the impacts of land management practices on wild bees outside of agricultural systems, including in forests managed intensively for wood production. We assessed changes in wild bee communities with time since harvest in 60 intensively managed Douglas-fir (Pseudotsuga menziesii) stands across a gradient in stand ages spanning a typical harvest rotation. We measured bee abundance, species richness, and alpha and beta diversity, as well as habitat characteristics (i.e., floral resources, nesting substrates, understory vegetation, and early seral forest in the surrounding landscape) during the spring and summer of 2018 and 2019. We found that bee abundance and species richness declined rapidly with stand age, decreasing by 61% and 48%, respectively, for every 5 years since timber harvest. Asymptotic estimates of Shannon and Simpson diversity were highest in stands 6-10 years post-harvest and lowest after the forest canopy had closed, ~11 years post-harvest. Bee communities in older stands were nested subsets of bee communities found in younger stands, indicating that changes were due to species loss rather than turnover as the stands aged. Bee abundance-but not species richness-was positively associated with floral resource density, and neither metric was associated with floral richness. The amount of early seral forest in the surrounding landscape seemed to enhance bee species richness in older, closed-canopy stands, but otherwise had little effect. Changes in the relative abundance of bee species did not relate to bee functional characteristics such as sociality, diet breadth, or nesting substrate. Our study demonstrates that Douglas-fir plantations develop diverse communities of wild bees shortly after harvest, but those communities erode rapidly over time as forest canopies close. Therefore, stand-scale management activities that prolong the precanopy closure period and enhance floral resources during the initial stage of stand regeneration will provide the greatest opportunity to enhance bee diversity in landscapes dominated by intensively managed conifer forests.

Fragmentation effects on an endangered species across a gradient from the interior to edge of its range.

Understanding how habitat fragmentation affects individual species is complicated by challenges associated with quantifying species-specific habitat and spatial variability in fragmentation effects within a species' range. We aggregated a 29-year breeding survey data set for the endangered marbled murrelet (Brachyramphus marmoratus) from >42,000 forest sites throughout the Pacific Northwest (Oregon, Washington, and northern California) of the United States. We built a species distribution model (SDM) in which occupied sites were linked with Landsat imagery to quantify murrelet-specific habitat and then used occupancy models to test the hypotheses that fragmentation negatively affects murrelet breeding distribution and that these effects are amplified with distance from the marine foraging habitat toward the edge of the species' nesting range. Murrelet habitat declined in the Pacific Northwest by 20% since 1988, whereas the proportion of habitat comprising edges increased by 17%, indicating increased fragmentation. Furthermore, fragmentation of murrelet habitat at landscape scales (within 2 km of survey stations) negatively affected occupancy of potential breeding sites, and these effects were amplified near the range edge. On the coast, the odds of occupancy decreased by 37% (95% confidence interval [CI] -54 to 12) for each 10% increase in edge habitat (i.e., fragmentation), but at the range edge (88 km inland) these odds decreased by 99% (95% CI 98 to 99). Conversely, odds of murrelet occupancy increased by 31% (95% CI 14 to 52) for each 10% increase in local edge habitat (within 100 m of survey stations). Avoidance of fragmentation at broad scales but use of locally fragmented habitat with reduced quality may help explain the lack of murrelet population recovery. Further, our results emphasize that fragmentation effects can be nuanced, scale dependent, and geographically variable. Awareness of these nuances is critical for developing landscape-level conservation strategies for species experiencing broad-scale habitat loss and fragmentation.

Efectos de la fragmentación sobre las especies en peligro a lo largo de un gradiente desde el interior hasta el borde de su distribución Resumen Es complicado entender el efecto de la fragmentación del hábitat sobre las especies individuales debido a los retos asociados con la cuantificación de hábitats específicos por especie y la variabilidad espacial de los efectos de la fragmentación dentro de la distribución de la especie. Combinamos los datos de un censo reproductivo realizado durante 29 años para el mérgulo jaspeado (Brachyramphus marmoratus) de >42,000 sitios boscosos a lo largo del noroeste del Pacífico (Oregón, Washington, y el norte de California, EE. UU.). Construimos un modelo de distribución de especie (MDE) en el cual los sitios ocupados estuvieron vinculados con imágenes de Landsat para cuantificar el hábitat específico del mérgulo y después usamos los modelos de ocupación para comprobar la hipótesis de que la fragmentación afecta negativamente la distribución reproductiva de la especie y que estos efectos se amplifican con la distancia entre el hábitat de forrajeo marino y el borde de la distribución de anidación de la especie. El hábitat del mérgulo declinó en la zona en un 20% a partir de 1988, mientras que la proporción de hábitat que comprende bordes incrementó en un 17%, lo que indica un aumento en la fragmentación. Además, la fragmentación del hábitat del mérgulo a escala de paisaje (a de 2 km de las estaciones de censo) afectó negativamente a la ocupación de sitios potenciales de reproducción y estos efectos se amplificaron cerca del borde de la distribución. La probabilidad de ocupación disminuyó en un 37% (95% IC -54 a 12) por cada 10% de incremento en el hábitat de borde (es decir, fragmentación) en la costa, pero en el borde de la distribución (88 km tierra adentro), esta probabilidad disminuyó en un 99% (95% IC 98 a 99). De forma contraria, la probabilidad de ocupación incrementó en un 31% (95% IC 14 a 52) por cada 10% de incremento en el hábitat de borde local (a 100 m de las estaciones de censo). La evasión de la fragmentación a gran escala y el uso de hábitats con calidad reducida y fragmentados a nivel local podría explicar la falta de recuperación poblacional del mérgulo. Más allá, nuestros resultados resaltan que los efectos de la fragmentación pueden estar matizados, depender de la escala y tener variación geográfica. Es importante tener conciencia de estos matices para desarrollar estrategias de conservación a nivel paisaje para las especies que experimentan fragmentación y pérdida del hábitat a gran escala.

Pollinator foraging tactics have divergent consequences for the mating system of a tropical plant.

Resolving the consequences of pollinator foraging behaviour for plant mating systems is a fundamental challenge in evolutionary ecology. Pollinators may adopt particular foraging tactics: complete trapline foraging (repeated movements along a fixed route), sample-and-shift trapline foraging (a variable route that incorporates information from previous experiences) and territorial foraging (stochastic movements within a restricted area). Studies that integrate these pollinator foraging tactics with plant mating systems are generally lacking. We investigate the consequences of particular pollinator foraging tactics for Heliconia tortuosa. We combine parentage and sibship inference analysis with simulation modelling to: estimate mating system parameters; infer the foraging tactic adopted by the pollinators; and quantify the impact of pollinator foraging tactics on mating system parameters. We found high outcrossing rates, ubiquitous multiple paternity and a pronounced departure from near-neighbour mating. We also found that plants repeatedly receive pollen from a series of particular donors. We infer that the pollinators primarily adopt complete trapline foraging and occasionally engage in sample-and-shift trapline foraging. This enhances multiple paternity without a substantial increase in near-neighbour mating. The particular pollinator foraging tactics have divergent consequences for multiple paternity and near-neighbour mating. Thus, pollinator foraging behaviour is an important driver of the ecology and evolution of plant mating systems.

Forest microclimate and composition mediate long-term trends of breeding bird populations.

Climate change is contributing to biodiversity redistributions and species declines. However, cooler microclimate conditions provided by old-growth forest structures compared with surrounding open or younger forests have been hypothesized to provide thermal refugia for species that are sensitive to climate warming and dampen the negative effects of warming on population trends of animals (i.e., the microclimate buffering hypothesis). In addition to thermal refugia, the compositional and structural diversity of old-growth forest vegetation itself may provide resources to species that are less available in forests with simpler structure (i.e., the insurance hypothesis). We used 8 years of breeding bird abundance data from a forested watershed, accompanied with sub-canopy temperature data, and ground- and LiDAR-based vegetation data to test these hypotheses and identify factors influencing bird population changes from 2011 to 2018. After accounting for imperfect detection, we found that for 5 of 20 bird species analyzed, abundance trends tended to be less negative or neutral at sites with cooler microclimates, which supports the microclimate buffering hypothesis. Negative effects of warming on two species were also reduced in locations with greater forest compositional diversity supporting the insurance hypothesis. We provide the first empirical evidence that complex forest structure and vegetation diversity confer microclimatic advantages to some animal populations in the face of climate change. Conservation of old-growth forests, or their characteristics in managed forests, could help slow the negative effects of climate warming on some breeding bird populations via microclimate buffering and possibly insurance effects.

Elevated inbreeding in Heliconia tortuosa is determined by tropical forest stand age, isolation and loss of hummingbird functional diversity.

Forest conversion and habitat loss are major threats to biological diversity. Forest regeneration can mitigate the negative effects of old-growth forest loss on species diversity, but less is known about the extent to which forest loss reduces genetic diversity in remnant populations and whether secondary forests play a role in the maintenance of genetic diversity. We quantified genetic diversity in a tropical hummingbird-pollinated understorey herb, Heliconia tortuosa, across a landscape mosaic of primary and secondary forest regrowth. Using microsatellite genotypes from >850 adult and juvenile plants within 33 forest patches and extensive bird surveys, we examined the effect of contemporary and historical landscape features including forest age (primary vs. secondary forest), stand isolation and pollinator assemblages on genetic diversity and levels of inbreeding in H. tortuosa. We found that inbreeding was up to three times higher in secondary forest, and this effect was amplified with reductions in primary forest in the surrounding landscape through reduced observed heterozygosity in isolated fragments. Inbreeding in forest patches was negatively correlated with the local frequency of specialist long-distance foraging traplining hummingbirds. Traplining hummingbirds therefore appear to facilitate mating among unrelated plants-an inference we tested using empirically parameterized simulations. Higher levels of inbreeding in H. tortuosa are therefore associated with reduced functional diversity of hummingbirds in secondary forests and forest patches isolated from primary forests. Our findings suggest a cryptic consequence of primary forest loss and secondary forest regeneration through the disruption of mutualistic interactions resulting in the erosion of genetic diversity in a common understorey plant.

Forest degradation drives widespread avian habitat and population declines.

In many regions of the world, forest management has reduced old forest and simplified forest structure and composition. We hypothesized that such forest degradation has resulted in long-term habitat loss for forest-associated bird species of eastern Canada (130,017 km2) which, in turn, has caused bird-population declines. Despite little change in overall forest cover, we found substantial reductions in old forest as a result of frequent clear-cutting and a broad-scale transformation to intensified forestry. Back-cast species distribution models revealed that breeding habitat loss occurred for 66% of the 54 most common species from 1985 to 2020 and was strongly associated with reduction in old age classes. Using a long-term, independent dataset, we found that habitat amount predicted population size for 94% of species, and habitat loss was associated with population declines for old-forest species. Forest degradation may therefore be a primary cause of biodiversity decline in managed forest landscapes.

Connectivity conservation at the crossroads: protected areas versus payments for ecosystem services in conserving connectivity for Colombian carnivores.

Protected areas (PAs) constitute one of the main tools for global landscape conservation. Recently, payments for environmental services (PES) have attracted interest from national and regional governments and are becoming one of the leading conservation policy instruments in tropical countries. However, the degree to which areas designated for PES overlap with areas that are critical for maintaining species' landscape connectivity is rarely evaluated. We estimated habitat distributions and connectivity for 16 of the 22 mammalian carnivores occurring in the Caribbean region of Colombia, and identified the overlap between existing PAs and areas identified as being important for connectivity for these species. We also evaluated the potential impact of creation of new PAs versus new PES areas on conserving connectivity for carnivores. Our results show that PAs cover only a minor percentage of the total area that is important for maintaining connectivity ( x = 26.8 % ± 20.2 s . d . ). On the other hand, PES, if implemented extensively, could contribute substantially to mammalian carnivores' connectivity ( x = 45.4 % ± 12.8 s . d . ). However, in a more realistic scenario with limited conservation investment in which fewer areas are set aside, a strategy based on implementing new PAs seems superior to PES. We argue that prioritizing designation of new PAs will be the most efficient means through which to maintain connectivity.

The macroecology of landscape ecology.

One of landscape ecology's main goals is to unveil how biodiversity is impacted by habitat transformation. However, the discipline suffers from significant context dependency in observed spatial and temporal trends, hindering progress towards understanding the mechanisms driving species declines and preventing the development of accurate estimates of future biodiversity change. Here, we discuss recent evidence that populations' and species' responses to habitat change at the landscape scale are modulated by factors and processes occurring at macroecological scales, such as historical disturbance rates, distance to geographic range edges, and climatic suitability. We suggest that placing landscape ecology studies in a macroecological lens will help to explain seemingly inconsistent results and will ultimately create better predictive models to help mitigate the biodiversity crisis.

The evolution, ecology, and conservation of hummingbirds and their interactions with flowering plants.

The ecological co-dependency between plants and hummingbirds is a classic example of a mutualistic interaction: hummingbirds rely on floral nectar to fuel their rapid metabolisms, and more than 7000 plant species rely on hummingbirds for pollination. However, threats to hummingbirds are mounting, with 10% of 366 species considered globally threatened and 60% in decline. Despite the important ecological implications of these population declines, no recent review has examined plant-hummingbird interactions in the wider context of their evolution, ecology, and conservation. To provide this overview, we (i) assess the extent to which plants and hummingbirds have coevolved over millions of years, (ii) examine the mechanisms underlying plant-hummingbird interaction frequencies and hummingbird specialization, (iii) explore the factors driving the decline of hummingbird populations, and (iv) map out directions for future research and conservation. We find that, despite close associations between plants and hummingbirds, acquiring evidence for coevolution (versus one-sided adaptation) is difficult because data on fitness outcomes for both partners are required. Thus, linking plant-hummingbird interactions to plant reproduction is not only a major avenue for future coevolutionary work, but also for studies of interaction networks, which rarely incorporate pollinator effectiveness. Nevertheless, over the past decade, a growing body of literature on plant-hummingbird networks suggests that hummingbirds form relationships with plants primarily based on overlapping phenologies and trait-matching between bill length and flower length. On the other hand, species-level specialization appears to depend primarily on local community context, such as hummingbird abundance and nectar availability. Finally, although hummingbirds are commonly viewed as resilient opportunists that thrive in brushy habitats, we find that range size and forest dependency are key predictors of hummingbird extinction risk. A critical direction for future research is to examine how potential stressors - such as habitat loss and fragmentation, climate change, and introduction of non-native plants - may interact to affect hummingbirds and the plants they pollinate.

Reconciling biodiversity with timber production and revenue via an intensive forest management experiment.

Understanding how land-management intensification shapes the relationships between biodiversity, yield, and economic benefit is critical for managing natural resources. Yet, manipulative experiments that test how herbicides affect these relationships are scarce, particularly in forest ecosystems where considerable time lags exist between harvest revenue and initial investments. We assessed these relationships by combining 7 yr of biodiversity surveys (>800 taxa) and forecasts of timber yield and economic return from a replicated, large-scale experiment that manipulated herbicide application intensity in operational timber plantations. Herbicides reduced species richness across trophic groups (-18%), but responses by higher-level trophic groups were more variable (0-38% reduction) than plant responses (-40%). Financial discounting, a conventional economic method to standardize past and future cash flows, strongly modified biodiversity-revenue relationships caused by management intensity. Despite a projected 28% timber yield gain with herbicides, biodiversity-revenue trade-offs were muted when opportunity costs were high (i.e., economic discount rates ≥7%). Although herbicides can drive biodiversity-yield trade-offs, under certain conditions, financial discounting provides opportunities to reconcile biodiversity conservation with revenue.

When are hypotheses useful in ecology and evolution?

Research hypotheses have been a cornerstone of science since before Galileo. Many have argued that hypotheses (1) encourage discovery of mechanisms, and (2) reduce bias-both features that should increase transferability and reproducibility. However, we are entering a new era of big data and highly predictive models where some argue the hypothesis is outmoded. We hypothesized that hypothesis use has declined in ecology and evolution since the 1990s, given the substantial advancement of tools further facilitating descriptive, correlative research. Alternatively, hypothesis use may have become more frequent due to the strong recommendation by some journals and funding agencies that submissions have hypothesis statements. Using a detailed literature analysis (N = 268 articles), we found prevalence of hypotheses in eco-evo research is very low (6.7%-26%) and static from 1990-2015, a pattern mirrored in an extensive literature search (N = 302,558 articles). Our literature review also indicates that neither grant success nor citation rates were related to the inclusion of hypotheses, which may provide disincentive for hypothesis formulation. Here, we review common justifications for avoiding hypotheses and present new arguments based on benefits to the individual researcher. We argue that stating multiple alternative hypotheses increases research clarity and precision, and is more likely to address the mechanisms for observed patterns in nature. Although hypotheses are not always necessary, we expect their continued and increased use will help our fields move toward greater understanding, reproducibility, prediction, and effective conservation of nature.

Producing wood at least cost to biodiversity: integrating Triad and sharing-sparing approaches to inform forest landscape management.

Forest loss and degradation are the greatest threats to biodiversity worldwide. Rising global wood demand threatens further damage to remaining native forests. Contrasting solutions across a continuum of options have been proposed, yet which of these offers most promise remains unresolved. Expansion of high-yielding tree plantations could free up forest land for conservation provided this is implemented in tandem with stronger policies for conserving native forests. Because plantations and other intensively managed forests often support far less biodiversity than native forests, a second approach argues for widespread adoption of extensive management, or 'ecological forestry', which better simulates natural forest structure and disturbance regimes - albeit with compromised wood yields and hence a need to harvest over a larger area. A third, hybrid suggestion involves 'Triad' zoning where the landscape is divided into three sorts of management (reserve, ecological/extensive management, and intensive plantation). Progress towards resolving which of these approaches holds the most promise has been hampered by the absence of a conceptual framework and of sufficient empirical data formally to identify the most appropriate landscape-scale proportions of reserves, extensive, and intensive management to minimize biodiversity impacts while meeting a given level of demand for wood. In this review, we argue that this central challenge for sustainable forestry is analogous to that facing food-production systems, and that the land sharing-sparing framework devised to establish which approach to farming could meet food demand at least cost to wild species can be readily adapted to assess contrasting forest management regimes. We develop this argument in four ways: (i) we set out the relevance of the sharing-sparing framework for forestry and explore the degree to which concepts from agriculture can translate to a forest management context; (ii) we make design recommendations for empirical research on sustainable forestry to enable application of the sharing-sparing framework; (iii) we present overarching hypotheses which such studies could test; and (iv) we discuss potential pitfalls and opportunities in conceptualizing landscape management through a sharing-sparing lens. The framework we propose will enable forest managers worldwide to assess trade-offs directly between conservation and wood production and to determine the mix of management approaches that best balances these (and other) competing objectives. The results will inform ecologically sustainable forest policy and management, reduce risks of local and global extinctions from forestry, and potentially improve a valuable sector's social license to operate.

A forest loss report card for the world's protected areas.

Protected areas are a key tool in the conservation of global biodiversity and carbon stores. We conducted a global test of the degree to which more than 18,000 terrestrial protected areas (totalling 5,293,217 km2) reduce deforestation in relation to unprotected areas. We also derived indices that quantify how well countries' forests are protected, both in terms of forested area protected and effectiveness of protected areas at reducing deforestation, in relation to vertebrate species richness, aboveground forest carbon biomass and background deforestation rates. Overall, protected areas did not eliminate deforestation, but reduced deforestation rates by 41%. Protected area deforestation rates were lowest in small reserves with low background deforestation rates. Critically, we found that after adjusting for effectiveness, only 6.5%-rather than 15.7%-of the world's forests are protected, well below the Aichi Convention on Biological Diversity's 2020 Target of 17%. We propose that global targets for protected areas should include quantitative goals for effectiveness in addition to spatial extent.

Do birds help trees grow? An experimental study of the effects of land-use intensification on avian trophic cascades.

By regulating populations of herbivores, predators can indirectly influence plant production. However, the factors influencing the strength of this type of trophic cascade are still unclear. We hypothesized that changes to plant community structure would affect the number of avian predators, thereby mediating cascade strength. Using a 4-yr, blocked, split-plot experiment, we independently manipulated both predators (birds) and plants in an early seral managed forest system in western Oregon, USA, and measured abundance across three trophic levels. We applied herbicides, as a surrogate for land-use intensification, to recently clear-cut stands to establish an experimental gradient in plant abundance and species richness, and excluded birds using 28, 225 m2 exclosures. In total, we counted and identified 94,738 arthropods of 141 families in paired control and bird exclosure plots. On average, insectivorous birds reduced arthropod abundance by 16% and plant damage by 14%, and some well-known pests (e.g., Adelges cooleyi) of crop trees (mostly Pseudotsuga menziesii) in our system were reduced by as much as 30%. However, this effect did not translate into a trophic cascade that increased crop-tree growth in the presence of birds. We experimentally reduced plant abundance and diversity by 67% and 55%, respectively, in the most intensive herbicide treatment in relation to untreated controls, but reduced vegetative resources did not change the strength of the direct effect of birds on arthropods or the indirect effect of birds on plants.