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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

A fire deficit persists across diverse North American forests despite recent increases in area burned

Rapid increases in wildfire area burned across North American forests pose novel challenges for managers and society. Increasing area burned raises questions about whether, and to what degree, contemporary fire regimes (1984–2022) are still departed from historical fire regimes (pre-1880). We use the North American tree-ring fire-scar network (NAFSN), a multi-century record comprising >1800 fire-scar sites spanning diverse forest types, and contemporary fire perimeters to ask whether there is a contemporary fire surplus or fire deficit, and whether recent fire years are unprecedented relative to historical fire regimes. Our results indicate, despite increasing area burned in recent decades, that a widespread fire deficit persists across a range of forest types and recent years with exceptionally high area burned are not unprecedented when considering the multi-century perspective offered by fire-scarred trees. For example, ‘record’ contemporary fire years such as 2020 burned 6% of NAFSN sites—the historical average—well below the historical maximum of 29% sites that burned in 1748. Although contemporary fire extent is not unprecedented across many North American forests, there is abundant evidence that unprecedented contemporary fire severity is driving forest loss in many ecosystems and adversely impacting human lives, infrastructure, and water supplies.

Nature Communications

Climate, environment, and disturbance history govern resilience of western North American Forests

Before the advent of intensive forest management and fire suppression, western North American forests exhibited a naturally occurring resistance and resilience to wildfires and other disturbances. Resilience, which encompasses resistance, reflects the amount of disruption an ecosystem can withstand before its structure or organization qualitatively shift to a different basin of attraction. In fire-maintained forests, resilience to disturbance events arose primarily from vegetation pattern-disturbance process interactions at several levels of organization. Using evidence from 15 ecoregions, spanning forests from Canada to Mexico, we review the properties of forests that reinforced qualities of resilience and resistance. We show examples of multi-level landscape resilience, of feedbacks within and among levels, and how conditions have changed under climatic and management influences. We highlight geographic similarities and important differences in the structure and organization of historical landscapes, their forest types, and in the conditions that have changed resilience and resistance to abrupt or large-scale disruptions. We discuss the role of the regional climate in episodically or abruptly reorganizing plant and animal biogeography and forest resilience and resistance to disturbances. We give clear examples of these changes and suggest that managing for resilient forests is a construct that strongly depends on scale and human social values. It involves human communities actively working with the ecosystems they depend on, and the processes that shape them, to adapt landscapes, species, and human communities to climate change while maintaining core ecosystem processes and services. Finally, it compels us to embrace management approaches that incorporate ongoing disturbances and anticipated effects of climatic changes, and to support dynamically shifting patchworks of forest and non-forest. Doing so could make these shifting forest conditions and wildfire regimes less disruptive to individuals and society.

western North America

Spatiotemporal synchrony of climate and fire occurrence across North American forests (1750-1880)

Aim Increasing aridity has driven widespread synchronous fire occurrence in recent decades across North America. The lack of historical (pre-1880) fire records limits our ability to understand long-term continental fire-climate dynamics. The goal of this study is to use tree-ring reconstructions to determine the relationships between spatiotemporal patterns in historical climate and widespread fire occurrence in North American forests, and whether they are stable through time. This information will address a major knowledge gap required to inform projections of future fire. Location North American Forests. Time Period 1750–1880 CE. Major Taxa Studies Trees. Methods We applied regionalisation methods to tree-ring reconstructions of historical summer soil moisture and annual fire occurrence to independently identify broad- and fine-scale climate and fire regions based on common inter-annual variability. We then tested whether the regions were stable through time and for spatial correspondence between the climate and fire regions. Last, we used correlation analysis to quantify the strength of the fire-climate associations through time. Results We found that broad-scale historical patterns in climate and fire have strong spatial coherence. Although climate and fire regions vary over time, large core areas of the regions were stable. The association between climate and fire varied through time and was strongest in western North America, likely due to a combination of factors, such as the magnitude of drought frequency and severity, as well as varying use of fire by human communities. Main Conclusions The historical perspective gained through tree-ring reconstructions of climate and fire patterns and their association suggests that climate-driven synchrony of fire across large areas of the continent in recent decades is not unprecedented, will likely continue into the future, and may exhibit similar spatial patterns.

North America

Selection and quality assessment of Landsat data for the North American forest dynamics forest history maps of the US

Using the NASA Earth Exchange platform, the North American Forest Dynamics (NAFD) project mapped forest history wall-to-wall, annually for the contiguous US (1986–2010) using the Vegetation Change Tracker algorithm. As with any effort to identify real changes in remotely sensed time-series, data gaps, shifts in seasonality, misregistration, inconsistent radiometry and cloud contamination can be sources of error. We discuss the NAFD image selection and processing stream (NISPS) that was designed to minimize these sources of error. The NISPS image quality assessments highlighted issues with the Landsat archive and metadata including inadequate georegistration, unreliability of the pre-2009 L5 cloud cover assessments algorithm, missing growing-season imagery and paucity of clear views. Assessment maps of Landsat 5–7 image quantities and qualities are presented that offer novel perspectives on the growing-season archive considered for this study. Over 150,000+ Landsat images were considered for the NAFD project. Optimally, one high quality cloud-free image in each year or a total of 12,152 images would be used. However, to accommodate data gaps and cloud/shadow contamination 23,338 images were needed. In 220 specific path-row image years no acceptable images were found resulting in data gaps in the annual national map products.

International Journal of Digital Earth

A case for using Plethodontid salamanders for monitoring biodiversity and ecosystem integrity of North American forests

Terrestrial salamanders of the family Plethodontidae have unique attributes that make them excellent indicators of biodiversity and ecosystem integrity in forested habitats. Their longevity, small territory size, site fidelity, sensitivity to natural and anthropogenic perturbations, tendency to occur in high densities, and low sampling costs mean that counts of plethodontid salamanders provide numerous advantages over counts of other North American forest organisms for indicating environmental change. Furthermore, they are tightly linked physiologically to microclimatic and successional processes that influence the distribution and abundance of numerous other hydrophilic but difficult-to-study forest-dwelling plants and animals. Ecosystem processes such as moisture cycling, food-web dynamics, and succession, with their related structural and microclimatic variability, all affect forest biodiversity and have been shown to affect salamander populations as well. We determined the variability associated with sampling for plethodontid salamanders by estimating the coefficient of variation (CV ) from available time-series data. The median coefficient of variation indicated that variation in counts of individuals among studies was much lower in plethodontids (27%) than in lepidoptera (93%), passerine birds (57%), small mammals (69%), or other amphibians (37–46%), which means plethodontid salamanders provide an important statistical advantage over other species for monitoring long-term forest health.

Conservation Biology

Range-wide salamander densities reveal a key component of terrestrial vertebrate biomass in eastern North American forests

Characterizing the population density of species is a central interest in ecology. Eastern North America is the global hotspot for biodiversity of plethodontid salamanders, an inconspicuous component of terrestrial vertebrate communities, and among the most widespread is the eastern red-backed salamander, Plethodon cinereus . Previous work suggests population densities are high with significant geographic variation, but comparisons among locations are challenged by lack of standardization of methods and failure to accommodate imperfect detection. We present results from a large-scale research network that accounts for detection uncertainty using systematic survey protocols and robust statistical models. We analysed mark–recapture data from 18 study areas across much of the species range. Estimated salamander densities ranged from 1950 to 34 300 salamanders ha −1 , with a median of 9965 salamanders ha −1 . We compared these results to previous estimates for P. cinereus and other abundant terrestrial vertebrates. We demonstrate that overall the biomass of P. cinereus , a secondary consumer, is of similar or greater magnitude to widespread primary consumers such as white-tailed deer ( Odocoileus virginianus ) and Peromyscus mice, and two to three orders of magnitude greater than common secondary consumer species. Our results add empirical evidence that P. cinereus , and amphibians in general, are an outsized component of terrestrial vertebrate communities in temperate ecosystems.

Biology Letters

Giving ecological meaning to satellite-derived fire severity metrics across North American forests

Satellite-derived spectral indices such as the relativized burn ratio (RBR) allow fire severity maps to be produced in a relatively straightforward manner across multiple fires and broad spatial extents. These indices often have strong relationships with field-based measurements of fire severity, thereby justifying their widespread use in management and science. However, satellite-derived spectral indices have been criticized because their non-standardized units render them difficult to interpret relative to on-the-ground fire effects. In this study, we built a Random Forest model describing a field-based measure of fire severity, the composite burn index (CBI), as a function of multiple spectral indices, a variable representing spatial variability in climate, and latitude. CBI data primarily representing forested vegetation from 263 fires (8075 plots) across the United States and Canada were used to build the model. Overall, the model performed well, with a cross-validated R 2 of 0.72, though there was spatial variability in model performance. The model we produced allows for the direct mapping of CBI, which is more interpretable compared to spectral indices. Moreover, because the model and all spectral explanatory variables were produced in Google Earth Engine, predicting and mapping of CBI can realistically be undertaken on hundreds to thousands of fires. We provide all necessary code to execute the model and produce maps of CBI in Earth Engine. This study and its products will be extremely useful to managers and scientists in North America who wish to map fire effects over large landscapes or regions.

Remote Sensing

Ecological consequences of mountain pine beetle outbreaks for wildlife in western North American forests

Mountain pine beetle ( Dendroctonus ponderosae ) (MPB) outbreaks are increasingly prevalent in western North America, causing considerable ecological change in pine ( Pinus spp.) forests with important implications for wildlife. We reviewed studies examining wildlife responses to MPB outbreaks and postoutbreak salvage logging to inform forest management and guide future research. Our review included 16 studies describing MPB outbreak relationships with 89 bird species and 6 studies describing relationships with 11 mammalian species, but no studies of reptiles or amphibians. We included studies that compared wildlife response metrics temporally (before versus after the outbreak) and spatially (across sites that varied in severity of outbreak) in relation to beetle outbreaks. Outbreaks ranged in size from 20,600 to ≥10 7 ha and studies occurred 1‐30 years after the peak MPB outbreak, but most studies were conducted over the short-term (i.e., ≤6 years after the peak of MPB-induced tree mortality). Birds were the only taxa studied frequently; however, high variability existed among those studies to allow many inferences, although some patterns were evident. Avian studies concluded that cavity-nesting species responded more favorably to beetle-killed forests than species with open-cup nests, and species nesting in the shrub layer favored outbreak forests compared with ground and open-cup canopy nesters that generally showed mixed relationships. Bark-drilling species as a group clearly demonstrated a positive short-term association with MPB epidemics compared with that of other foraging assemblages. Cavity-nesting birds that do not consume bark beetles (i.e., secondary cavity-nesting species and nonbark-drilling woodpeckers) also exhibited some positive responses to MPB outbreaks, although not as pronounced or consistent as those of bark-drilling woodpeckers. Mammalian responses to MPB outbreaks were mixed. Studies consistently reported negative effects of MPB outbreaks on red squirrels ( Tamiasciurus hudsonicus ). However, there is evidence that red squirrels can persist after an outbreak under some conditions, e.g., when nonhost tree species are present. For small mammal species associated with forest understories, responses may be most pronounced during the postepidemic period (>6 years after the peak of beetle-induced tree mortality) when snags fall to produce coarse woody debris. Postoutbreak salvage logging studies ( n = 6) reported results that lacked consensus. Postoutbreak salvage logging may have an impact on fewer wildlife species than postfire salvage logging, probably because only host-specific tree species are removed after beetle outbreaks.

Forest Science

Improved fire severity mapping in the North American boreal forest using a hybrid composite method

Fire severity is a key driver shaping the ecological structure and function of North American boreal ecosystems, a biome dominated by large, high-intensity wildfires. Satellite-derived burn severity maps have been an important tool in these remote landscapes for both fire and resource management. The conventional methodology to produce satellite-inferred fire severity maps generally involves comparing imagery from 1 year before and 1 year after a fire, yet environmental conditions unique to the boreal have limited the accuracy of resulting products. We introduce an alternative method – the ‘hybrid composite’ – based on deriving mean severity over time on a per-pixel basis within the cloud-computing environment of Google Earth Engine. It constructs the post-fire image from satellite data composited from all valid images (i.e., clear-sky and snow-free) acquired in the time period immediately after fire through the early growing season of the following year. We compare this approach to paired-scene and composite approaches where the post-fire time period is from the growing season 1 year after fire. Validation statistics based on field-derived data for 52 fires across Alaska and Canada indicate that the hybrid composite method outperforms the other approaches. This approach presents an efficient and cost-effective means to monitor and explore trends and patterns across broad spatial domains, and could be applied to fires in other regions, especially those with frequent cloud cover or rapid vegetation recovery.

Alaska

Temporal bird community dynamics are strongly affected by landscape fragmentation in a Central American tropical forest region

Habitat loss and fragmentation are considered the main causes of species extinctions, particularly in tropical ecosystems. The objective of this work was to evaluate the temporal dynamics of tropical bird communities in landscapes with different levels of fragmentation in eastern Guatemala. We evaluated five bird community dynamic parameters for forest specialists and generalists: (1) species extinction, (2) species turnover, (3) number of colonizing species, (4) relative species richness, and (5) a homogeneity index. For each of 24 landscapes, community dynamic parameters were estimated from bird point count data, for the 1998–1999 and 2008–2009 periods, accounting for species’ detection probability. Forest specialists had higher extinction rates and a smaller number of colonizing species in landscapes with higher fragmentation, thus having lower species richness in both time periods. Alternatively, forest generalists elicited a completely different pattern, showing a curvilinear association to forest fragmentation for most parameters. Thus, greater community dynamism for forest generalists was shown in landscapes with intermediate levels of fragmentation. Our study supports general theory regarding the expected negative effects of habitat loss and fragmentation on the temporal dynamics of biotic communities, particularly for forest specialists, providing strong evidence from understudied tropical bird communities.

Biodiversity and Conservation

Vulnerability of carbon storage in North American boreal forests to wildfires during the 21st century

The boreal forest contains large reserves of carbon. Across this region, wildfires influence the temporal and spatial dynamics of carbon storage. In this study, we estimate fire emissions and changes in carbon storage for boreal North America over the 21st century. We use a gridded data set developed with a multivariate adaptive regression spline approach to determine how area burned varies each year with changing climatic and fuel moisture conditions. We apply the process‐based Terrestrial Ecosystem Model to evaluate the role of future fire on the carbon dynamics of boreal North America in the context of changing atmospheric carbon dioxide (CO 2 ) concentration and climate in the A2 and B2 emissions scenarios of the CGCM2 global climate model. Relative to the last decade of the 20th century, decadal total carbon emissions from fire increase by 2.5–4.4 times by 2091–2100, depending on the climate scenario and assumptions about CO 2 fertilization. Larger fire emissions occur with warmer climates or if CO 2 fertilization is assumed to occur. Despite the increases in fire emissions, our simulations indicate that boreal North America will be a carbon sink over the 21st century if CO 2 fertilization is assumed to occur in the future. In contrast, simulations excluding CO 2 fertilization over the same period indicate that the region will change to a carbon source to the atmosphere, with the source being 2.1 times greater under the warmer A2 scenario than the B2 scenario. To improve estimates of wildfire on terrestrial carbon dynamics in boreal North America, future studies should incorporate the role of dynamic vegetation to represent more accurately post‐fire successional processes, incorporate fire severity parameters that change in time and space, account for human influences through increased fire suppression, and integrate the role of other disturbances and their interactions with future fire regime.

Global Change Biology

Fire and drought

Historical and presettlement relationships between drought and wildfire have been well documented in much of North America, with forest fire occurrence and area burned clearly increasing in response to drought. Drought interacts with other controls (forest productivity, topography, and fire weather) to affect fire intensity and severity. Fire regime characteristics (area, frequency, severity) are the product of many individual fires, so both weather and climate - including short- and long-term droughts - are important. It is worth noting, however, that the factors controlling fire events and fire regimes are complex and extend beyond drought and climate alone, and so fire regimes and wildfires are affected by other variables from local-to-global scales. Fire history evidence from diverse climate regimes and forest ecosystems suggests that North American forest fire regimes were moderately to strongly controlled by climate prior to Euro-American settlement and subsequent fire exclusion and fire suppression (Flatley and others 2013, Hessl and others 2004, Heyerdahl and others 2002, Heyerdahl and others 2008, Swetnam 1990, Swetnam and Betancourt 1998, Weisberg and Swanson 2003). These presettlement fire histories indicate a relationship between low precipitation anomalies and widespread fire activity, especially in the Western United States. This is consistent with a regional depletion of soil and atmospheric moisture, which leads to low moisture in foliage and surface fuels and ultimately to the potential for widespread fire (Swetnam and Betancourt 1998). Some fire histories in the American Southwest also demonstrate a lagged relationship with above-average antecedent precipitation (Swetnam and Betancourt 1998) and/or cooler temperatures (Veblen and others 2000) in the year(s) prior to years of widespread fire. Most of these records are derived from fire-scarred trees that survived fire events and thus are primarily indicative of low- or mixedseverity fire regimes, although some work has focused also on evidence from high-severity fire regimes (Heyerdahl and others 2002).

Book chapter

Northern forest winters have lost cold, snowy conditions that are important for ecosystems and human communities

Winter is an understudied but key period for the socio-ecological systems of northeastern North American forests. A growing awareness of the importance of the winter season to forest ecosystems and surrounding communities has inspired several decades of research, both across the northern forest and at other mid- and high-latitude ecosystems around the globe. Despite these efforts, we lack a synthetic understanding of how winter climate change may impact hydrological and biogeochemical processes and the social and economic activities they support. Here we take advantage of 100 years of meteorological observations across the northern forest region of the northeastern U.S. and eastern Canada to develop a suite of indicators that enable a cross-cutting understanding of 1) how winter temperatures and snow cover have been changing and 2) how these shifts may impact both ecosystems and surrounding human communities. We show that cold and snow-covered conditions have generally decreased over the past 100 years. These trends suggest positive outcomes for tree health as related to reduced fine root mortality and nutrient loss associated with winter frost but negative outcomes as related to the northward advancement and proliferation of forest insect pests. In addition to effects on vegetation, reductions in cold temperatures and snow cover are likely to have negative impacts on the ecology of the northern forest through impacts on water, soils, and wildlife. The overall loss of coldness and snow cover may also have negative consequences for logging and forest products, vector-borne diseases and human health, recreation and tourism, and cultural practices, which together represent important social and economic dimensions for the northern forest region. These findings advance our understanding of how our changing winters may transform the socio- ecological system of a region that has been defined by the contrasting rhythm of the seasons. Our research also identifies a trajectory of change that informs our expectations for the future as the climate continues to warm.

Ecological Applications

Centennial-scale reductions in nitrogen availability in temperate forests of the United States

Forests cover 30% of the terrestrial Earth surface and are a major component of the global carbon (C) cycle. Humans have doubled the amount of global reactive nitrogen (N), increasing deposition of N onto forests worldwide. However, other global changes—especially climate change and elevated atmospheric carbon dioxide concentrations—are increasing demand for N, the element limiting primary productivity in temperate forests, which could be reducing N availability. To determine the long-term, integrated effects of global changes on forest N cycling, we measured stable N isotopes in wood, a proxy for N supply relative to demand, on large spatial and temporal scales across the continental U.S.A. Here, we show that forest N availability has generally declined across much of the U.S. since at least 1850 C.E. with cool, wet forests demonstrating the greatest declines. Across sites, recent trajectories of N availability were independent of recent atmospheric N deposition rates, implying a minor role for modern N deposition on the trajectory of N status of North American forests. Our results demonstrate that current trends of global changes are likely to be consistent with forest oligotrophication into the foreseeable future, further constraining forest C fixation and potentially storage.

Scientific Reports

The North American tree-ring fire-scar network

Fire regimes in North American forests are diverse and modern fire records are often too short to capture important patterns, trends, feedbacks, and drivers of variability. Tree-ring fire scars provide valuable perspectives on fire regimes, including centuries-long records of fire year, season, frequency, severity, and size. Here, we introduce the newly compiled North American tree-ring fire-scar network (NAFSN), which contains 2562 sites, >37,000 fire-scarred trees, and covers large parts of North America. We investigate the NAFSN in terms of geography, sample depth, vegetation, topography, climate, and human land use. Fire scars are found in most ecoregions, from boreal forests in northern Alaska and Canada to subtropical forests in southern Florida and Mexico. The network includes 91 tree species, but is dominated by gymnosperms in the genus Pinus . Fire scars are found from sea level to >4000-m elevation and across a range of topographic settings that vary by ecoregion. Multiple regions are densely sampled (e.g., >1000 fire-scarred trees), enabling new spatial analyses such as reconstructions of area burned. To demonstrate the potential of the network, we compared the climate space of the NAFSN to those of modern fires and forests; the NAFSN spans a climate space largely representative of the forested areas in North America, with notable gaps in warmer tropical climates. Modern fires are burning in similar climate spaces as historical fires, but disproportionately in warmer regions compared to the historical record, possibly related to under-sampling of warm subtropical forests or supporting observations of changing fire regimes. The historical influence of Indigenous and non-Indigenous human land use on fire regimes varies in space and time. A 20th century fire deficit associated with human activities is evident in many regions, yet fire regimes characterized by frequent surface fires are still active in some areas (e.g., Mexico and the southeastern United States). These analyses provide a foundation and framework for future studies using the hundreds of thousands of annually- to sub-annually-resolved tree-ring records of fire spanning centuries, which will further advance our understanding of the interactions among fire, climate, topography, vegetation, and humans across North America.

Ecosphere

Decreased atmospheric nitrogen deposition in eastern North America: Predicted responses of forest ecosystems

Historical increases in emissions and atmospheric deposition of oxidized and reduced nitrogen (N) provided the impetus for extensive, global-scale research investigating the effects of excess N in terrestrial and aquatic ecosystems, with several regions within the Eastern Deciduous Forest of the United States found to be susceptible to negative effects of excess N. The Clean Air Act and associated rules have led to decreases in emissions and deposition of oxidized N, especially in eastern U.S., representing a research challenge and opportunity for ecosystem ecologists and biogeochemists. The purpose of this paper is to predict changes in the structure and function of North American forest ecosystems in a future of decreased N deposition. Hysteresis is a property of a system wherein output is not a strict function of corresponding input, incorporating lag, delay, or history dependence, particularly when the response to decreasing input is different from the response to increasing input. We suggest a conceptual hysteretic model predicting varying lag times in recovery of soil acidification, plant biodiversity, soil microbial communities, forest carbon (C) and N cycling, and surface water chemistry toward pre-N impact conditions. Nearly all of these can potentially respond strongly to reductions in N deposition. Most responses are expected to show some degree of hysteresis, with the greatest delays in response occurring in processes most tightly linked to “slow pools” of N in wood and soil organic matter. Because experimental studies of declines in N loads in forests of North America are lacking and because of the expected hysteresis, it is difficult to generalize from experimental results to patterns expected from declining N deposition. These will likely be long-term phenomena, difficult to distinguish from other, concurrent environmental changes, including elevated atmospheric CO 2 , climate change, reductions in acidity, invasions of new species, and long-term vegetation responses to past disturbance.

Environmental Pollution

Disturbance-mediated accelerated succession in two Michigan forest types

In northern lower Michigan, logging accelerated sugar maple ( Acer saccharum ) dominance in a northern white cedar ( Thuja occidentals ) community, and clear-cutting and burning quickly converted certain sites dominated by mature jack pine ( Pinus banksiana ) to early-succesional hardwoods, including Prunus, Populus , and Quercus . In both forest types the succeeding hardwoods should continue to increase in the future at the expense of the pioneer conifer species. In the cedar example, sugar maple was also increasing a an undisturbed, old-growth stand, but at a much reduced rate than in the logged stand. Traditionally, disturbance was through to set back succession to some earlier stage. However, out study sites and at least several other North American forest communities exhibited accelerated succession following a wide range of disturbances, including logging fire, ice storms, wind-throw, disease, insect attack, and herbicide spraying.

Michigan