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.

Herbicides and herbivory interact to drive plant community and crop-tree establishment.

Land management practices often directly alter vegetation structure and composition, but the degree to which ecological processes such as herbivory interact with management to influence biodiversity is less well understood. We hypothesized that large herbivores compound the effects of intensive forest management on early seral plant communities and plantation establishment (i.e., tree survival and growth), and the degree of such effects is dependent on the intensity of management practices. We established 225-m2 wild-ungulate (deer and elk) exclosures, nested within a manipulated gradient of management intensity (no-herbicide control, light herbicide, moderate herbicide, and intensive herbicide treatments), replicated at the scale of whole harvest units (10-19 ha). Vegetation structure, composition, and crop-tree responses to herbivory varied across the gradient of herbicide application during the first two years of stand establishment, with herbivory effects most evident at light and moderate herbicide treatments. In the moderate herbicide treatment, which approximates management applied to >2.5 million hectares in the Pacific Northwest, United States, foraging by deer and elk resulted in simplified, low-cover plant communities more closely resembling the intensive herbicide treatment. Herbivory further suppressed the growth of competing vegetation in the light herbicide treatment, improving crop-tree survival, and providing early evidence of an ecosystem service. By changing community composition and vegetation structure, intensive forest management alters foraging selectivity and subsequent plant-herbivore interactions; initial shifts in early seral communities are likely to influence understory plant communities and tree growth in later stages of forest development.

Plant diversity enhances moth diversity in an intensive forest management experiment.

Intensive forest management (IFM) promises to help satisfy increasing global demand for wood but may come at the cost of local reductions to forest biodiversity. IFM often reduces early seral plant diversity as a result of efforts to eliminate plant competition with crop trees. If diversity is a function of bottom-up drivers, theory predicts that specialists at lower trophic levels (e.g., insect herbivores) should be particularly sensitive to reductions in plant diversity. We conducted a stand-level experiment to test bottom-up controls on moth community structure, as mediated by degrees of forest management intensity. Using a dataset of 12,003 moths representing 316 moth species, moth richness decreased only slightly, if at all, as herbicide intensity increased (P = 0.062); the moderate treatment, which is most commonly applied in the northwestern USA, was estimated to have 4.72 (±2.14 SE, P = 0.039) fewer species than the control. Structural equation modeling revealed strong support for an effect of herbicide on plant abundance, which influenced plant species richness and subsequently moth species richness. Moth species richness was associated with plant species richness and followed a power law function (z = 0.42, P = 0.006), which is surprisingly consistent with a recent large-scale experiment in agricultural systems, and provides support for bottom-up drivers of moth community structure. Moth abundance was not influenced by the direct effects of silvicultural herbicide treatments. Site-level effects and variation in pre-harvest vegetation communities resulted in residual broadleaf and herbaceous vegetation in even the most intensive treatment. Even at low densities, these residual deciduous and herbaceous plants supported higher than expected moth abundance and richness. We conclude that forest management practices that retain early seral vegetation diversity are the most likely to conserve moth communities.

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.

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.