The forests of the midwestern United States at Euro-American settlement: Spatial and physical structure based on contemporaneous survey data.

We present gridded 8 km-resolution data products of the estimated stem density, basal area, and biomass of tree taxa at Euro-American settlement of the midwestern United States during the middle to late 19th century for the states of Minnesota, Wisconsin, Michigan, Illinois, and Indiana. The data come from settlement-era Public Land Survey (PLS) data (ca. 0.8-km resolution) of trees recorded by land surveyors. The surveyor notes have been transcribed, cleaned, and processed to estimate stem density, basal area, and biomass at individual points. The point-level data are aggregated within 8 km grid cells and smoothed using a generalized additive statistical model that accounts for zero-inflated continuous data and provides approximate Bayesian uncertainty estimates. The statistical modeling smooths out sharp spatial features (likely arising from statistical noise) within areas smaller than about 200 km2. Based on this modeling, presettlement Midwestern landscapes supported multiple dominant species, vegetation types, forest types, and ecological formations. The prairies, oak savannas, and forests each had distinctive structures and spatial distributions across the domain. Forest structure varied from savanna (averaging 27 Mg/ha biomass) to northern hardwood (104 Mg/ha) and mesic southern forests (211 Mg/ha). The presettlement forests were neither unbroken and massively-statured nor dominated by young forests constantly structured by broad-scale disturbances such as fire, drought, insect outbreaks, or hurricanes. Most forests were structurally between modern second growth and old growth. We expect the data product to be useful as a baseline for investigating how forest ecosystems have changed in response to the last several centuries of climate change and intensive Euro-American land use and as a calibration dataset for paleoecological proxy-based reconstructions of forest composition and structure for earlier time periods. The data products (including raw and smoothed estimates at the 8-km scale) are available at the LTER Network Data Portal as version 1.0.

Evaluating a new method for reconstructing forest conditions from General Land Office survey records.

Historical forest conditions are often used to inform contemporary management goals because historical forests are considered to be resilient to ecological disturbances. The General Land Office (GLO) surveys of the late 19th and early 20th centuries provide regionally quasi-contiguous data sets of historical forests across much of the Western United States. Multiple methods exist for estimating tree density from point-based sampling such as the GLO surveys, including distance-based and area-based approaches. Area-based approaches have been applied in California mixed-conifer forests but their estimates have not been validated. To assess the accuracy and precision of plotless density estimators with potential for application to GLO data in this region, we imposed a GLO sampling scheme on six mapped forest stands of known densities (159-784 trees/ha) in the Sierra Nevada in California, USA, and Baja California Norte, Mexico. We compared three distance-based plotless density estimators (Cottam, Pollard, and Morisita) as well as two Voronoi area (VA) estimators, the Delincé and mean harmonic Voronoi density (MHVD), to the true densities. We simulated sampling schemes of increasing intensity to assess sampling error. The relative error (RE) of density estimates for the GLO sampling scheme ranged from 0.36 to 4.78. The least biased estimate of tree density in every stand was obtained with the Morisita estimator and the most biased was obtained with the MHVD estimator. The MHVD estimates of tree density were 1.2-3.8 times larger than the true densities and performed best in stands subject to fire exclusion for 100 yr. The Delincé approach obtained accurate estimates of density, implying that the Voronoi approach is theoretically sound but that its application in the MHVD was flawed. The misapplication was attributed to two causes: (1) the use of a crown scaling factor that does not correct for the number of trees sampled and (2) the persistent underestimate of the true VA due to a weak relationship between tree size and VA. The magnitude of differences between true densities and MHVD estimates suggest caution in using results based on the MHVD to inform management and restoration practices in the conifer forests of the American West.

Correction: Statistically-Estimated Tree Composition for the Northeastern United States at Euro-American Settlement.

[This corrects the article DOI: 10.1371/journal.pone.0150087.].

Novel and Lost Forests in the Upper Midwestern United States, from New Estimates of Settlement-Era Composition, Stem Density, and Biomass.

BACKGROUND: EuroAmerican land-use and its legacies have transformed forest structure and composition across the United States (US). More accurate reconstructions of historical states are critical to understanding the processes governing past, current, and future forest dynamics. Here we present new gridded (8x8km) reconstructions of pre-settlement (1800s) forest composition and structure from the upper Midwestern US (Minnesota, Wisconsin, and most of Michigan), using 19th Century Public Land Survey System (PLSS), with estimates of relative composition, above-ground biomass, stem density, and basal area for 28 tree types. This mapping is more robust than past efforts, using spatially varying correction factors to accommodate sampling design, azimuthal censoring, and biases in tree selection. CHANGES IN FOREST STRUCTURE: We compare pre-settlement to modern forests using US Forest Service Forest Inventory and Analysis (FIA) data to show the prevalence of lost forests (pre-settlement forests with no current analog), and novel forests (modern forests with no past analogs). Differences between pre-settlement and modern forests are spatially structured owing to differences in land-use impacts and accompanying ecological responses. Modern forests are more homogeneous, and ecotonal gradients are more diffuse today than in the past. Novel forest assemblages represent 28% of all FIA cells, and 28% of pre-settlement forests no longer exist in a modern context. Lost forests include tamarack forests in northeastern Minnesota, hemlock and cedar dominated forests in north-central Wisconsin and along the Upper Peninsula of Michigan, and elm, oak, basswood and ironwood forests along the forest-prairie boundary in south central Minnesota and eastern Wisconsin. Novel FIA forest assemblages are distributed evenly across the region, but novelty shows a strong relationship to spatial distance from remnant forests in the upper Midwest, with novelty predicted at between 20 to 60km from remnants, depending on historical forest type. The spatial relationships between remnant and novel forests, shifts in ecotone structure and the loss of historic forest types point to significant challenges for land managers if landscape restoration is a priority. The spatial signals of novelty and ecological change also point to potential challenges in using modern spatial distributions of species and communities and their relationship to underlying geophysical and climatic attributes in understanding potential responses to changing climate. The signal of human settlement on modern forests is broad, spatially varying and acts to homogenize modern forests relative to their historic counterparts, with significant implications for future management.

Statistically-Estimated Tree Composition for the Northeastern United States at Euro-American Settlement.

We present a gridded 8 km-resolution data product of the estimated composition of tree taxa at the time of Euro-American settlement of the northeastern United States and the statistical methodology used to produce the product from trees recorded by land surveyors. Composition is defined as the proportion of stems larger than approximately 20 cm diameter at breast height for 22 tree taxa, generally at the genus level. The data come from settlement-era public survey records that are transcribed and then aggregated spatially, giving count data. The domain is divided into two regions, eastern (Maine to Ohio) and midwestern (Indiana to Minnesota). Public Land Survey point data in the midwestern region (ca. 0.8-km resolution) are aggregated to a regular 8 km grid, while data in the eastern region, from Town Proprietor Surveys, are aggregated at the township level in irregularly-shaped local administrative units. The product is based on a Bayesian statistical model fit to the count data that estimates composition on the 8 km grid across the entire domain. The statistical model is designed to handle data from both the regular grid and the irregularly-shaped townships and allows us to estimate composition at locations with no data and to smooth over noise caused by limited counts in locations with data. Critically, the model also allows us to quantify uncertainty in our composition estimates, making the product suitable for applications employing data assimilation. We expect this data product to be useful for understanding the state of vegetation in the northeastern United States prior to large-scale Euro-American settlement. In addition to specific regional questions, the data product can also serve as a baseline against which to investigate how forests and ecosystems change after intensive settlement. The data product is being made available at the NIS data portal as version 1.0.

Four centuries of change in northeastern United States forests.

The northeastern United States is a predominately-forested region that, like most of the eastern U.S., has undergone a 400-year history of intense logging, land clearance for agriculture, and natural reforestation. This setting affords the opportunity to address a major ecological question: How similar are today's forests to those existing prior to European colonization? Working throughout a nine-state region spanning Maine to Pennsylvania, we assembled a comprehensive database of archival land-survey records describing the forests at the time of European colonization. We compared these records to modern forest inventory data and described: (1) the magnitude and attributes of forest compositional change, (2) the geography of change, and (3) the relationships between change and environmental factors and historical land use. We found that with few exceptions, notably the American chestnut, the same taxa that made up the pre-colonial forest still comprise the forest today, despite ample opportunities for species invasion and loss. Nonetheless, there have been dramatic shifts in the relative abundance of forest taxa. The magnitude of change is spatially clustered at local scales (<125 km) but exhibits little evidence of regional-scale gradients. Compositional change is most strongly associated with the historical extent of agricultural clearing. Throughout the region, there has been a broad ecological shift away from late successional taxa, such as beech and hemlock, in favor of early- and mid-successional taxa, such as red maple and poplar. Additionally, the modern forest composition is more homogeneous and less coupled to local climatic controls.