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Shading decreases the abundance of the herbivorous California horn snail, Cerithidea californica

Most of the intertidal zone in estuaries of California, USA and Baja California, Mexico is covered with vascular vegetation. Shading by these vascular plants influences abiotic and biotic processes that shape benthic community assemblages. We present data on the effects of shading on the California horn snail, Cerithidea californica . This species is important because it is the most common benthic macrofaunal species in these systems and acts as an obligate intermediate host of several species of rematode parasites that infect several other species. Using observational and experimental studies, we found a negative effect of shade on the distribution and abundance of the California horn snail. We hypothesized that shading reduces the abundance of the epipelic diatoms that the snails feeds on, causing snails to leave haded areas. We observed a negative relationship between vascular plant cover, sub-canopy light levels, and snail density in Mugu Lagoon. Then we experimentally manipulated light regimes, by clipping vegetation and adding shade structures, and found higher snail densities at higher light levels. In Goleta Slough, we isolated the effect of shade from vegetation by documenting a negative relationship between the shade created by two bridges and diatom and snail densities. We also found that snails moved the greatest distances over shaded channel banks compared to unshaded channel banks. Further, we documented the effect of water depth and channel bank orientation on shading in this system. An additional effect of shading is the reduction of temperature, providing an alternative explanation for some of our results. These results broaden our knowledge of how variation in the light environment influences the ecology of estuarine ecosystems.

California↗

Effectiveness of environmental flows for riparian restoration in arid regions: A tale of four rivers

Environmental flows have become important restoration tools on regulated rivers. However, environmental flows are often constrained by other demands within the river system and thus typically are comprised of smaller water volumes than the natural flows they are meant to replace, which can limit their functional efficacy. We review environmental flow programs aimed at restoring riparian vegetation on four arid zone rivers: the Tarim River in China; the Bill Williams River in Arizona, U.S.; the delta of the Colorado River in Mexico; and the Murrumbidgee River in southern Australia. Our goal is to determine what worked and what did not work to accomplish restoration goals. The lower Tarim River in China formerly formed a “green corridor” across the Taklamakan Desert. The riparian zone deteriorated due to diversion of surface and groundwater for irrigated agriculture. A massive restoration program began in 2000 with release of 1038 million cubic meters of water over the first three years. Groundwater levels rose but the ecological response was less than expected politically, socially and within the scientific community. However, releases continued and by 2015 portions of the original iconic Populus euphratica (Euphrates poplar) forest were reestablished. The natural flow regime of the Bill Williams River was disrupted by construction of a dam in 1968, dramatically reducing peak flows along with associated fluvial processes. As a result, the channel narrowed and riparian vegetation expanded and was comprised largely of an introduced shrub species ( Tamarix spp.). Environmental flow releases including small, managed floods and sustained base flows have been implemented since the mid 1990’s to promote establishment and maintenance of native riparian trees (cottonwoods and willows) and have been successful, although in a “downsized” portion of the valley bottom. Experience from the Bill Williams was used to help design the Minute 319 environmental flow in the delta of the Colorado River in 2014. Water was released as a short, one-time pulse during spring with the intent of starting new cohorts of cottonwood and willow. However, fluvial disturbance was limited by the relatively small magnitude pulse, low flows did not continue throughout the growing season in some reaches, native tree recruitment was low, and most of the new plants recruited were Tamarix . The inundated portion of the floodplain did respond with a temporary increase in greenness as measured by satellite vegetation indices, however. The Murrumbidgee River in Australia is a tributary in the Murray-Darling River Basin, which supports iconic red gum ( Eucalyptus camaldulensis ) forests that depend on near-yearly floods for maintenance. During the recent Millennial Drought (2000–2010) environmental flows were provided on an experimental basis to small portions of the Yanga National Forest to see how much water was needed. As with the Colorado River delta, gains in vegetation vigor as measured by satellite vegetation indices following the flows were temporary. Environmental flows in the Bill Williams were able to restore enough overbank flooding and fluvial disturbance to promote some establishment of new cohorts of trees, but on the Colorado and Murrumbidgee Rivers larger volumes of total flows released over longer periods and targeted restoration will be needed to restore the ecosystems. A measure of success in restoring the Euphrates poplar forest on the Tarim and germinating new chorts of willows on the Bill Williams has been achieved after 15–20 years of environmental flows, but the Colorado River delta and Murrumbidgee Rivers have only received one or two flows. Success in enhancing native trees in the Colorado delta has been achieved in restoration plots, but the Murrumbidgee will require large overbank flows on a continuing schedule to rejuvenate the red gum forest.

Ecological Engineering↗

Estimating the aboveground biomass and carbon stocks of tall shrubs in a prerestoration degraded salt marsh

Wetlands play a vital role in Earth's carbon cycle and provide important ecosystem services. Their ability to perform their roles can be compromised by human activities that destroy or impair their functioning. The restoration of degraded wetlands may allow carbon cycle functioning, as well as other services, to be recovered. Predicting the potential outcomes from any restoration project requires upfront consideration, including via modeling possible changes in carbon stocks. In this study, we quantified the carbon stocks in tall shrub vegetation proliferating in a degraded salt marsh that is currently the subject of an extensive restoration project. We produced allometric models to estimate biomass and carbon stocks for three tall shrub species, which, along with other freshwater and upland species in the area, will die with continued restoration. Therefore, estimating the potential for carbon losses in biomass is important. We also developed a means of estimating carbon stocks in other nontree plants in the estuary area. Useful equations for estimating the biomass of tall shrubs are limited in general and lacking for degraded systems. Our study adds to the literature on carbon stocks in shrub species and fills a data gap for degraded ecosystems. It also contributes to the broader carbon feasibility study of the aforementioned restoration project that was designed to predict the overall net impact of the project on greenhouse gas emissions in the ecosystem.

Restoration Ecology↗

Prospective HyspIRI global observations of tidal wetlands

Tidal wetlands are highly productive and act as critical habitat for a wide variety of plants, fish, shellfish, and other wildlife. These ecotones between aquatic and terrestrial environments also provide protection from storm damage, run-off filtering, and recharge of aquifers. Many wetlands along coasts have been exposed to stress-inducing alterations globally, including dredge and fill operations, hydrologic modifications, pollutants, impoundments, fragmentation by roads/ditches, and sea level rise. For wetland protection and sensible coastal development, there is a need to monitor these ecosystems at global and regional scales. Recent advances in satellite sensor design and data analysis are providing practical methods for monitoring natural and man-made changes in wetlands. However, available satellite remote sensors have been limited to mapping primarily wetland location and extent. This paper describes how the HyspIRI hyperspectral and thermal infrared sensors can be used to study and map key ecological properties, such as species composition, biomass, hydrology, and evapotranspiration of tidal salt and brackish marshes and mangroves, and perhaps other major wetland types, including freshwater marshes and wooded/shrub wetlands.

Remote Sensing of Environment↗

Nesting ecology and reproductive rate of the red-winged blackbird in tidal marshes of the upper Chesapeake Bay region

The nesting ecology and reproductive rate of the polygynous red-winged blackbird, Agelaius phoeniceus, were studied in the tidal marshes of Maryland during the period of 1958 through 1961. Sixteen nesting colonies were located in six major marsh communities of the Eastern Shore and were visited approximately twice a week from late April to mid-August. The average clutch size for 537 clutches was 3.3 eggs, with a range of 2 to 5 eggs. The ratio of territorial males to nesting females was 1:1.9. There was direct evidence of double broods by four females. The average number of young produced was 4.2 per breeding female or 8.1 per breeding male. Nesting success for the total of 675 active nests was 57%, with a range from 38% to 69% in the colonies. Robust plants that held constant form throughout the nesting season supported 95% of the nests, and the success of these nests was 58%; in contrast, non-robust plants supported only 5% of the nests, and success of this group of nests was only 26%. Nesting success varied with height from ground: 45% for nests less than 2 feet from the ground, 55% for those 2 to 4 feet, and 62% for those more than 4 feet. Histories of 749 nests were summarized by stages: 749 newly built, 675 with eggs, 424 with nestlings, and 388 with fledging young.

Chesapeake Bay region↗

Ecology of red swamps in the glaciated northeast: a community profile

This report is part of a series of profiles on the ecology of wetland and deepwater habitats. This particular profile addresses red maple swamps in the glaciated northeastern United States. Red maple (Acer rubrum) swamp is a dominant wetland type in most of the region; it reaches its greatest abundance in southern New England and northern New Jersey, where it comprises 60-800/o of all inland wetlands. Red maple swamps occur in a wide variety of hydrogeologic settings, from small, isolated basins in till or glaciofluvial deposits to extensive wetland complexes on glacial lake beds, and from hillside seeps to stream floodplains and lake edges. Individual swamps may be seasonally flooded, temporarily flooded, or seasonally saturated, and soils may be mineral or organic. As many as five distinct vegetation layers may occur in these swamps, including trees, saplings, shrubs, herbs, and ground cover plants such as bryophytes and clubmosses. On a regional scale, red maple swamps support at least 50 species of trees, more than 90 species of shrubs and vines, and more than 300 species of nonwoody plants. These swamps also provide habitat for a rich faunal community, including several wetland-dependent species. In areas that are becoming urbanized, these wetlands often constitute critical habitat for facultative species as well. Red maple swamps also are important sites for flood storage, water quality improvement, recreation, scenic beauty, and open space.

Connecticut;Maine;Massachusetts;New Hampshire;New ↗

Are migratory waterfowl vectors of seagrass pathogens?

Migratory waterfowl vector plant seeds and other tissues, but little attention has focused on the potential of avian vectoring of plant pathogens. Extensive meadows of eelgrass (Zostera marina) in southwest Alaska support hundreds of thousands of waterfowl during fall migration and may be susceptible to plant pathogens. We recovered DNA of organisms pathogenic to eelgrass from environmental samples and in the cloacal contents of eight of nine waterfowl species that annually migrate along the Pacific coast of North America and Asia. Coupled with a signal of asymmetrical gene flow of eelgrass running counter to that expected from oceanic and coastal currents between Large Marine Ecosystems, this evidence suggests waterfowl are vectors of eelgrass pathogens.

Alaska↗

Climate-induced tree-mortality pulses are obscured by broad-scale and long-term greening

Vegetation greening has been suggested to be a dominant trend over recent decades, but severe pulses of tree mortality in forests after droughts and heatwaves have also been extensively reported. These observations raise the question of to what extent the observed severe pulses of tree mortality induced by climate could affect overall vegetation greenness across spatial grains and temporal extents. To address this issue, here we analyse three satellite-based datasets of detrended growing-season normalized difference vegetation index (NDVI GS ) with spatial resolutions ranging from 30 m to 8 km for 1,303 field-documented sites experiencing severe drought- or heat-induced tree-mortality events around the globe. We find that severe tree-mortality events have distinctive but localized imprints on vegetation greenness over annual timescales, which are obscured by broad-scale and long-term greening. Specifically, although anomalies in NDVI GS (ΔNDVI) are negative during tree-mortality years, this reduction diminishes at coarser spatial resolutions (that is, 250 m and 8 km). Notably, tree-mortality-induced reductions in NDVI GS (|ΔNDVI|) at 30-m resolution are negatively related to native plant species richness and forest height, whereas topographic heterogeneity is the major factor affecting ΔNDVI differences across various spatial grain sizes. Over time periods of a decade or longer, greening consistently dominates all spatial resolutions. The findings underscore the fundamental importance of spatio-temporal scales for cohesively understanding the effects of climate change on forest productivity and tree mortality under both gradual and abrupt changes.

Nature Ecology & Evolution↗

The human dimension of fire regimes on Earth

Humans and their ancestors are unique in being a fire‐making species, but ‘natural’ (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human‐set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from ‘natural’ background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke‐related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas‐induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre‐human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Journal of Biogeography↗

Modeling impacts of drought-induced salinity intrusion on carbon dynamics in tidal freshwater forested wetlands

Tidal freshwater forested wetlands (TFFW) provide critical ecosystem services including essential habitat for a variety of wildlife species and significant carbon sinks for atmospheric carbon dioxide. However, large uncertainties remain concerning the impacts of climate change on the magnitude and variability of carbon fluxes and storage across a range of TFFW. In this study, we developed a process-driven Tidal Freshwater Wetlands DeNitrification-DeComposition model (TFW-DNDC) that has integrated new features, such as soil salinity effects on plant productivity and soil organic matter decomposition to explore carbon dynamics in TFFW in response to drought-induced saltwater intrusion. Eight sites along the floodplains of the Waccamaw River (USA) and the Savannah River (USA) were selected to represent TFFW transition from healthy to moderately and highly salt-impacted forests, and eventually to oligohaline marshes. TFW-DNDC was calibrated and validated using field observed annual litterfall, stem growth, root growth, soil heterotrophic respiration and soil organic carbon storage. Analyses indicate that plant productivity and soil carbon sequestration in TFFW could change substantially in response to increased soil porewater salinity and reduced soil water table due to drought, but in interactive ways dependent on the river simulated. Such responses are variable due to non-linear relationships between carbon cycling processes and environmental drivers. Plant productivity, plant respiration, soil organic carbon sequestration rate and storage in the highly salt-impacted forest sites decreased significantly under drought conditions compared to normal conditions. Considering the high likelihood of healthy and moderately salt-impacted forests becoming highly salt-impacted forests under future climate change and sea-level rise, it is very likely that TFFW will lose their capacity as carbon sinks without up-slope migration.

Georgia, South Carolina↗

Multiple models guide strategies for agricultural nutrient reductions

In response to degraded water quality, federal policy makers in the US and Canada called for a 40% reduction in phosphorus (P) loads to Lake Erie, and state and provincial policy makers in the Great Lakes region set a load-reduction target for the year 2025. Here, we configured five separate SWAT (US Department of Agriculture's Soil and Water Assessment Tool) models to assess load reduction strategies for the agriculturally dominated Maumee River watershed, the largest P source contributing to toxic algal blooms in Lake Erie. Although several potential pathways may achieve the target loads, our results show that any successful pathway will require large-scale implementation of multiple practices. For example, one successful pathway involved targeting 50% of row cropland that has the highest P loss in the watershed with a combination of three practices: subsurface application of P fertilizers, planting cereal rye as a winter cover crop, and installing buffer strips. Achieving these levels of implementation will require local, state/provincial, and federal agencies to collaborate with the private sector to set shared implementation goals and to demand innovation and honest assessments of water quality-related programs, policies, and partnerships.

Frontiers in Ecology and the Environment↗

Acute salt marsh dieback in the Mississippi River deltaic plain: A drought-induced phenomenon?

Aims Extensive dieback of salt marsh dominated by the perennial grass Spartina alterniflora occurred throughout the Mississippi River deltaic plain during 2000. More than 100,000 ha were affected, with 43,000 ha severely damaged. The aim of this work was to determine if sudden dieback could have been caused by a coincident drought and to assess the significance of this event with respect to long-term changes in coastal vegetation. Location Multiple dieback sites and reference sites were established along 150 km of shoreline in coastal Louisiana, USA. Methods Aerial and ground surveys were conducted from June 2000 to September 2001 to assess soil conditions and plant mortality and recovery. Results Dieback areas ranged in size from???300 m2-5 km2 in area with 50-100% mortality of plant shoots and rhizomes in affected zones. Co-occurring species such as Avicennia germinans (black mangrove) and Juncus roemerianus (needlegrass rush) were unaffected. Historical records indicate that precipitation, river discharge, and mean sea level were unusually low during the previous year. Although the cause of dieback is currently unknown, plant and soil characteristics were consistent with temporary soil desiccation that may have reduced water availability, increased soil salinity, and/or caused soil acidification (via pyrite oxidation) and increased uptake of toxic metals such as Fe or Al. Plant recovery 15 months after dieback was variable (0-58% live cover), but recovering plants were vigorous and indicated no longlasting effects of the dieback agent. Main conclusions These findings have relevance for global change models of coastal ecosystems that predict vegetation responses based primarily on long-term increases in sea level and submergence of marshes. Our results suggest that large-scale changes in coastal vegetation may occur over a relatively short time span through climatic extremes acting in concert with sea-level fluctuations and pre-existing soil conditions. ?? 2004 Blackwell Publishing Ltd.

Global Ecology and Biogeography↗

Physical, Chemical, Ecological, and Age Data and Trench Logs from Surficial Deposits at Hatch Point, Southeastern Utah

This report presents data and describes the methodology for physical, chemical and ecological measurements of sediment, soil, and vegetation, as well as age determinations of surficial deposits at Hatch Point, Canyon Rims area, Colorado Plateau, southeastern Utah. The results presented in this report support a study that examines geomorphic and soil factors that may influence boundaries between shrubland and grassland ecosystems in the study area. Shrubland ecosystems dominated by sagebrush (Artemisia tridentata) and grassland ecosystems dominated by native perennial grasses (for example, Hilaria jamesii and Sporabolis sp.) are high-priority conservation targets for the Federal Bureau of Land Management (BLM) and other resource managers because of their diversity, productivity, and vital importance as wildlife habitat. These ecosystems have been recognized as imperiled on a regional scale since at least the mid-1990s due to habitat loss (type conversions), land-use practices, and invasive exotic plants. In the Intermountain West, the exotic annual cheatgrass (Bromus tectorum) is recognized as one of the most pervasive and serious threats to the health of native sagebrush and grassland ecosystems through effects on fire regimes and resource conditions experienced by native species.

Open-File Report↗

A tour de force by Hawaii’s invasive mammals: Establishment, takeover, and ecosystem restoration through eradication

Invasive mammals have irreversibly altered ecosystems of Hawai‘i and other tropical Pacific islands in numerous cases through novel herbivory, predation, and diseases, thereby causing the disproportionate extinction of flora and fauna that occur nowhere else on Earth. The control and eradication of invasive mammals is the single most expensive management activity necessary for restoring ecological integrity to many natural areas of Hawai‘i and other Pacific Islands, and have already advanced the restoration of native biota by removing herbivorous ungulates from >750 km 2 . Rodenticides which have been tested and registered for hand and aerial broadcast in Hawai‘i have been used to eradicate rats from remote islands to protect nesting seabirds and are now being applied to larger islands to protect forest birds. The exclusion of other invasive mammals is now being undertaken with more sophisticated control techniques and fences. New fence designs are capable of excluding all mammals from areas to protect endangered native birds. Although the eradication of mammals from large areas has resulted in the restoration of some ecosystem processes such as natural forest regeneration, changes in other processes such as fire regimes, nutrient cycling, and invasive plant proliferation remain more difficult to reverse at larger landscape scales.

Hawaii↗

Spatial models can improve the experimental design of field-based transplant gardens by preventing bias due to neighborhood crowding

Field-based transplant gardens, including common and reciprocal garden experiments, are a powerful tool for studying genetic variation and gene-by-environment interactions. These experiments assume that individuals within the garden represent independent replicates growing in a homogenous environment. Plant neighborhood interactions are pervasive across plant populations and could violate assumptions of transplant garden experiments. We demonstrate how spatially explicit models for plant–plant interactions can provide novel insights on genotypes' performance in field-transplant garden designs. We used individual-based models, based on data from a sagebrush ( Artemisia spp.) common garden, to simulate the impact of spatial plant–plant interactions on between-group differences in plant growth. We found that planting densities within the range of those used in many common gardens can bias experimental outcomes. Our results demonstrate that higher planting densities can lead to inflated group differences and may confound genotypes' competitive ability and genetically underpinned variation. Synthesis. We propose that spatially explicit models can help avoid biased results by informing the design and analysis of field-based transplant garden experiments. Alternately, including neighborhood effects in post hoc analyses of transplant garden experiments is likely to provide novel insights into the roles of biotic factors and density dependence in genetic differentiation.

Ecology and Evolution↗

Nutrient fluxes at the landscape level and the R* rule

Nutrient cycling in terrestrial ecosystems involves not only the vertical recycling of nutrients at specific locations in space, but also biologically driven horizontal fluxes between different areas of the landscape. This latter process can result in net accumulation of nutrients in some places and net losses in others. We examined the effects of such nutrient-concentrating fluxes on the R * rule, which predicts that the species that can survive in steady state at the lowest level of limiting resource, R *, can exclude all competing species. To study the R * rule in this context, we used a literature model of plant growth and nutrient cycling in which both nutrients and light may limit growth, with plants allocating carbon and nutrients between foliage and roots according to different strategies. We incorporated the assumption that biological processes may concentrate nutrients in some parts of the landscape. We assumed further that these processes draw nutrients from outside the zone of local recycling at a rate proportional to the local biomass density. Analysis showed that at sites where there is a sufficient biomass-dependent accumulation of nutrients, the plant species with the highest biomass production rates (roughly corresponding to the best competitors) do not reduce locally available nutrients to a minimum concentration level (that is, minimum R *), as expected from the R * rule, but instead maximize local nutrient concentration. These new results require broadening of our understanding of the relationships between nutrients and vegetation competition on the landscape level. The R * rule is replaced by a more complex criterion that varies across a landscape and reduces to the R * rule only under certain limiting conditions.

Ecological Modelling↗

Demography with drones: Detecting growth and survival of shrubs with unoccupied aerial systems

Large-scale disturbances, such as megafires, motivate restoration at equally large extents. Measuring the survival and growth of individual plants plays a key role in current efforts to monitor restoration success. However, the scale of modern restoration (e.g., >10,000 ha) challenges measurements of demographic rates with field data. In this study, we demonstrate how unoccupied aerial system (UAS) flights can provide an efficient solution to the tradeoff of precision and spatial extent in detecting demographic rates from the air. We flew two, sequential UAS flights at two sagebrush ( Artemisia tridentata ) common gardens to measure the survival and growth of individual plants. The accuracy of Bayesian-optimized segmentation of individual shrub canopies was high (73–95%, depending on the year and site), and remotely sensed survival estimates were within 10% of ground-truthed survival censuses. Stand age structure affected remotely sensed estimates of growth; growth was overestimated relative to field-based estimates by 57% in the first garden with older stands, but agreement was high in the second garden with younger stands. Further, younger stands (similar to those just after disturbance) with shorter, smaller plants were sometimes confused with other shrub species and bunchgrasses, demonstrating a need for integrating spectral classification approaches that are increasingly available on affordable UAS platforms. The older stand had several merged canopies, which led to an underestimation of abundance but did not bias remotely sensed survival estimates. Advances in segmentation and UAS structure from motion photogrammetry will enable demographic rate measurements at management-relevant extents.

Restoration Ecology↗

Emerging themes in the ecology and management of North American forests

The 7th North American Forest Ecology Workshop, consisting of 149 presentations in 16 oral sessions and a poster session, reflected a broad range of topical areas currently under investigation in forest ecology and management. There was an overarching emphasis on the role of disturbance, both natural and anthropogenic, in the dynamics of forest ecosystems, and the recognition that legacies from past disturbances strongly influence future trajectories. Climate was invoked as a major driver of ecosystem change. An emphasis was placed on application of research findings for predicting system responses to changing forest management initiatives. Several “needs” emerged from the discussions regarding approaches to the study of forest ecosystems, including (1) consideration of variable spatial and temporal scales, (2) long-term monitoring, (3) development of universal databases more encompassing of time and space to facilitate meta-analyses, (4) combining field studies and modeling approaches, (5) standardizing methods of measurement and assessment, (6) guarding against oversimplification or overgeneralization from limited site-specific results, (7) greater emphasis on plant-animal interactions, and (8) better alignment of needs and communication of results between researchers and managers.

International Journal of Forestry Research↗