Native American geography shaped historical fire frequency in forests of eighteenth-century Pennsylvania, USA.

Researchers have debated the relative importance of environmental versus Indigenous effects on past fire regimes in eastern North America. Tree-ring fire-scar records (FSRs) provide local-resolution physical evidence of past fire, but few studies have spatially correlated fire frequency from FSRs with environmental and anthropogenic variables. No study has compared FSR locations to Native American settlement features in the eastern United States. We assess whether FSRs in the eastern US are located near regions of past Native American settlement. We also assess relationships between distance to Native American settlement, environmental conditions, and fire frequency in central Pennsylvania (PA), US, using an "ensemble of small models" approach for low sample sizes. Regression models of fire frequency at 21 locations in central PA often selected distance-based proxies of Indigenous land use. Models with mean annual temperature and Native American variables as predictors explained > 70% of the variation in fire frequency. Alongside temperature and wind speed, "distance to nearest trail" and "mean distance to nearest town" were significant and important predictors. In 18th-century central PA, fires were more frequent near Indigenous trails and towns, and further south due to increasing temperature and pyrophilic vegetation. However, for the entire eastern US, FSRs are located far from past settlement, limiting their effectiveness in detecting fire patterns near population centers. Improving understanding of historical fire will require developing FSRs closer to past Native American settlement.

Re-framing deer herbivory as a natural disturbance regime with ecological and socioeconomic outcomes in the eastern United States.

Natural disturbances are critical ecosystem processes, with both ecological and socioeconomic benefits and disadvantages. Large herbivores are natural disturbances that have removed plant biomass for millions of years, although herbivore influence likely has declined during the past thousands of years corresponding with extinctions and declines in distributions and abundances of most animal species. Nonetheless, the conventional view, particularly in eastern North America, is that herbivory by large wild herbivores is at unprecedented levels, resulting in unnatural damage to forests. Here, we propose consideration of large herbivores as a natural disturbance that also imparts many crucial ecological advantages, using white-tailed deer (Odocoileus virginianus), the only wild large herbivore remaining throughout the eastern U.S., as our focal species. We examined evidence of detrimental effects of browsing on trees and forbs. We then considered that deer contribute to both fuel reduction and ecological restoration of herbaceous plants and historical open forests of savannas and woodlands by controlling tree and shrub densities, mimicking the consumer role of fire. Similarly to other disturbances, deer disturbance 'regimes' are uneven in severity across different ecosystems and landscapes, resulting in heterogeneity and diversity. In addition to biodiversity support and fuel reduction, socioeconomic benefits include >$20 billion dollars per year by 10 million hunters that support jobs and wildlife agencies, non-consumptive enjoyment of nature by 80 million people, cultural importance, and deer as ecological ambassadors, whereas costs include about $5 billion and up to 450 human deaths per year for motor vehicle accidents, along with crop damage and disease transmission. From a perspective of historical ecology rather than current baselines, deer impart a fundamental disturbance process with many ecological benefits and a range of socioeconomic effects.

Addressing regional relationships between white-tailed deer densities and land classes.

White-tailed deer (Odocoileus virginianus) populations have recovered to about 30 million animals in the United States, but land cover has changed during the interval of recovery. To address the relationship between deer densities and current land cover at regional scales, I applied random forests and extreme gradient boosting classifiers to model low and high deer density classes, at two different thresholds (5.8 and 11.6 deer/km2), and land classes in three regions during approximately 2003. For low and high deer density classes divided at 5.8 deer/km2, deciduous broadleaf forest overall was the most influential and positive variable in the central east and central regions and crop and pasture were the most influential and negative variables in the southeast region. Deer density increased with area of deciduous and mixed forests, woody wetlands, and shrub in all regions. Deer density decreased with area of crop, developed open space, and developed low and medium residential density in all regions. For density classes divided at 11.6 deer/km2, deer density had the strongest relationship with woody wetlands in the central east region, mixed and deciduous forest in the southeast region, and woody wetlands and herbaceous vegetation in the central region. Deer density increased with deciduous and mixed forests, woody wetlands, and shrub in all regions. Conversely, deer density decreased with herbaceous vegetation, crop, and developed low residential densities in all regions. Therefore, at regional scales, deer overall occurred at greater densities in forests and woody wetlands and lower densities in agricultural and residential development, which did not appear to support more deer. Deer preference for forests does result in damage to forest products, but alternatively, some may consider that deer provide important socioeconomic and ecological services by reducing number of small trees, particularly in the absence of other disturbances that historically controlled tree biomass.

Future forest aboveground carbon dynamics in the central United States: the importance of forest demographic processes.

The Central Hardwood Forest (CHF) in the United States is currently a major carbon sink, there are uncertainties in how long the current carbon sink will persist and if the CHF will eventually become a carbon source. We used a multi-model ensemble to investigate aboveground carbon density of the CHF from 2010 to 2300 under current climate. Simulations were done using one representative model for each of the simple, intermediate, and complex demographic approaches (ED2, LANDIS PRO, and LINKAGES, respectively). All approaches agreed that the current carbon sink would persist at least to 2100. However, carbon dynamics after current carbon sink diminishes to zero differ for different demographic modelling approaches. Both the simple and the complex demographic approaches predicted prolonged periods of relatively stable carbon densities after 2100, with minor declines, until the end of simulations in 2300. In contrast, the intermediate demographic approach predicted the CHF would become a carbon source between 2110 and 2260, followed by another carbon sink period. The disagreement between these patterns can be partly explained by differences in the capacity of models to simulate gross growth (both birth and subsequent growth) and mortality of short-lived, relatively shade-intolerant tree species.

The history of widespread decrease in oak dominance exemplified in a grassland-forest landscape.

Regionally-distinctive open oak forest ecosystems have been replaced either by intensive agriculture and grazing fields or by denser forests throughout eastern North America and Europe. To quantify changes in tree communities and density in the Missouri Plains, a grassland-forest landscape, we used historical surveys from 1815 to 1864 and current surveys from 2004 to 2008. To estimate density for historical communities, we used the Morisita plotless density estimator and applied corrections for surveyor bias. To estimate density for current forests, we used Random Forests, an ensemble regression tree method, to predict densities from known values at plots using terrain and soil predictors. Oak species decreased from 62% of historical composition to 30% of current composition and black and white oaks historically were dominant species across 93% of the landscape and currently were dominant species across 42% of the landscape. Current forest density was approximately two times greater than historical densities, demonstrating loss of savanna and woodlands and transition to dense forest structure. Average tree diameters were smaller than in the past, but mean basal area and stocking remained similar over time because of the increase in density in current forests. Nevertheless, there were spatial differences; basal area and stocking decreased along rivers and increased away from rivers. Oak species are being replaced by other species in the Missouri Plains, similar to replacement throughout the range of Quercus. Long-term commitment to combinations of prescribed burning and silvicultural prescriptions in more xeric sites may be necessary for oak recruitment. Restoration of open oak ecosystems is a time-sensitive issue because restoration will become increasingly costly as oaks are lost from the overstory and the surrounding matrix becomes dominated by non-oak species.

Winning and losing tree species of reassembly in Minnesota's mixed and broadleaf forests.

We examined reassembly of winning and losing tree species, species traits including shade and fire tolerance, and associated disturbance filters and forest ecosystem types due to rapid forest change in the Great Lakes region since 1850. We identified winning and losing species by changes in composition, distribution, and site factors between historical and current surveys in Minnesota's mixed and broadleaf forests. In the Laurentian Mixed Forest, shade-intolerant aspen replaced shade-intolerant tamarack as the most dominant tree species. Fire-tolerant white pine and jack pine decreased, whereas shade-tolerant ashes, maples, and white cedar increased. In the Eastern Broadleaf Forest, fire-tolerant white oaks and red oaks decreased, while shade-tolerant ashes, American basswood, and maples increased. Tamarack, pines, and oaks have become restricted to sites with either wetter or sandier and drier soils due to increases in aspen and shade-tolerant, fire-sensitive species on mesic sites. The proportion of shade-tolerant species increased in both regions, but selective harvest reduced the applicability of functional groups alone to specify winners and losers. Harvest and existing forestry practices supported aspen dominance in mixed forests, although without aspen forestry and with fire suppression, mixed forests will transition to a greater composition of shade-tolerant species, converging to forests similar to broadleaf forests. A functional group framework provided a perspective of winning and losing species and traits, selective filters, and forest ecosystems that can be generalized to other regions, regardless of species identity.

Pseudoabsence generation strategies for species distribution models.

BACKGROUND: Species distribution models require selection of species, study extent and spatial unit, statistical methods, variables, and assessment metrics. If absence data are not available, another important consideration is pseudoabsence generation. Different strategies for pseudoabsence generation can produce varying spatial representation of species. METHODOLOGY: We considered model outcomes from four different strategies for generating pseudoabsences. We generating pseudoabsences randomly by 1) selection from the entire study extent, 2) a two-step process of selection first from the entire study extent, followed by selection for pseudoabsences from areas with predicted probability <25%, 3) selection from plots surveyed without detection of species presence, 4) a two-step process of selection first for pseudoabsences from plots surveyed without detection of species presence, followed by selection for pseudoabsences from the areas with predicted probability <25%. We used Random Forests as our statistical method and sixteen predictor variables to model tree species with at least 150 records from Forest Inventory and Analysis surveys in the Laurentian Mixed Forest province of Minnesota. CONCLUSIONS: Pseudoabsence generation strategy completely affected the area predicted as present for species distribution models and may be one of the most influential determinants of models. All the pseudoabsence strategies produced mean AUC values of at least 0.87. More importantly than accuracy metrics, the two-step strategies over-predicted species presence, due to too much environmental distance between the pseudoabsences and recorded presences, whereas models based on random pseudoabsences under-predicted species presence, due to too little environmental distance between the pseudoabsences and recorded presences. Models using pseudoabsences from surveyed plots produced a balance between areas with high and low predicted probabilities and the strongest relationship between density and area with predicted probabilities ≥75%. Because of imperfect accuracy assessment, the best assessment currently may be evaluation of whether the species has been sufficiently but not excessively predicted to occur.

Regime shifts and weakened environmental gradients in open oak and pine ecosystems.

Fire suppression allows tree species that are intolerant of fire stress to increase their distribution, potentially resulting in disruption of historical species-environmental relationships. To measure changes between historical General Land Office surveys (1815 to 1850) and current USDA Forest Inventory and Assessment surveys (2004 to 2008), we compared composition, distribution, and site factors of 21 tree species or species groups in the Missouri Ozarks. We used 24 environmental variables and random forests as a classification method to model distributions. Eastern redcedar, elms, maples, and other fire-sensitive species have increased in dominance in oak forests, with concurrent reductions by oak species; specific changes varied by ecological subsection. Ordinations displayed loss of separation between formerly distinctive oak and fire-sensitive tree species groups. Distribution maps showed decreased presence of disturbance-dependent oak and pine species and increased presence of fire-sensitive species that generally expanded from subsections protected from fire along rivers to upland areas, except for eastern redcedar, which expanded into these subsections. Large scale differences in spatial gradients between past and present communities paralleled reduced influence of local topographic gradients in the varied relief of the Missouri Ozarks, as fire-sensitive species have moved to higher, drier, and sunnier sites away from riverine corridors. Due to changes in land use, landscapes in the Missouri Ozarks, eastern United States, and world-wide are changing from open oak and pine-dominated ecosystems to novel oak-mixed species forests, although at fine scales, forests are becoming more diverse in tree species today. Fire suppression weakened the influence by environmental gradients over species dominance, allowing succession from disturbance-dependent oaks to an alternative state of fire-sensitive species. Current and future research and conservation that rely on historical relationships and ecological principles based on disturbance across the landscape will need to incorporate modern interactions among species for resources into management plans and projections.