Papers by David Mladenoff

SciVee, Feb 29, 2008
Human land use of forested regions has intensified worldwide in recent decades, threatening long-... more Human land use of forested regions has intensified worldwide in recent decades, threatening long-term sustainability. Primary effects include conversion of land cover or reversion to an earlier stage of successional development. Both types of change can have cascading effects through ecosystems; however, the longterm effects where forests are allowed to regrow are poorly understood. We quantify the regionalscale consequences of a century of Euro-American land use in the northern U.S. Great Lakes region using a combination of historical Public Land Survey records and current forest inventory and land cover data. Our analysis shows a distinct and rapid trajectory of vegetation change toward historically unprecedented and simplified conditions. In addition to overall loss of forestland, current forests are marked by lower species diversity, functional diversity, and structural complexity compared to pre-Euro-American forests. Today's forest is marked by dominance of broadleaf deciduous species-all 55 ecoregions that comprise the region exhibit a lower relative dominance of conifers in comparison to the pre-Euro-American period. Aspen (Populus grandidentata and P. tremuloides) and maple (Acer saccharum and A. rubrum) species comprise the primary deciduous species that have replaced conifers. These changes reflect the cumulative effects of local forest alterations over the region and they affect future ecosystem conditions as well as the ecosystem services they provide.

Proceedings of the National Academy of Sciences of the United States of America, Apr 14, 2009
One-third of net CO2 emissions to the atmosphere since 1850 are the result of land-use change, pr... more One-third of net CO2 emissions to the atmosphere since 1850 are the result of land-use change, primarily from the clearing of forests for timber and agriculture, but quantifying these changes is complicated by the lack of historical data on both former ecosystem conditions and the extent and spatial configuration of subsequent land use. Using fine-resolution historical survey records, we reconstruct pre-EuroAmerican settlement (1850s) forest carbon in the state of Wisconsin, examine changes in carbon after logging and agricultural conversion, and assess the potential for future sequestration through forest recovery. Results suggest that total aboveground live forest carbon (AGC) fell from 434 TgC before settlement to 120 TgC at the peak of agricultural clearing in the 1930s and has since recovered to approximately 276 TgC. The spatial distribution of AGC, however, has shifted significantly. Former savanna ecosystems in the south now store more AGC because of fire suppression and forest ingrowth, despite the fact that most of the region remains in agriculture, whereas northern forests still store much less carbon than before settlement. Across the state, continued sequestration in existing forests has the potential to contribute an additional 69 TgC. Reforestation of agricultural lands, in particular, the formerly high C-density forests in the north-central region that are now agricultural lands less optimal than those in the south, could contribute 150 TgC. Restoring historical carbon stocks across the landscape will therefore require reassessing overall land-use choices, but a range of options can be ranked and considered under changing needs for ecosystem services. forest-agriculture trade-offs ͉ old-growth forest ͉ Eastern North America ͉ land-use history ͉ carbon sink
Ecology and Recovery of Eastern Old-Growth Forests, 2018

PLOS ONE, 2021
We present gridded 8 km-resolution data products of the estimated stem density, basal area, and b... more 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 do...

Proceedings of the National Academy of Sciences, 2009
One-third of net CO 2 emissions to the atmosphere since 1850 are the result of land-use change, p... more One-third of net CO 2 emissions to the atmosphere since 1850 are the result of land-use change, primarily from the clearing of forests for timber and agriculture, but quantifying these changes is complicated by the lack of historical data on both former ecosystem conditions and the extent and spatial configuration of subsequent land use. Using fine-resolution historical survey records, we reconstruct pre-EuroAmerican settlement (1850s) forest carbon in the state of Wisconsin, examine changes in carbon after logging and agricultural conversion, and assess the potential for future sequestration through forest recovery. Results suggest that total above-ground live forest carbon (AGC) fell from 434 TgC before settlement to 120 TgC at the peak of agricultural clearing in the 1930s and has since recovered to approximately 276 TgC. The spatial distribution of AGC, however, has shifted significantly. Former savanna ecosystems in the south now store more AGC because of fire suppression and f...

Oikos, 2003
Self-organization describes the evolution process of complex structures where systems emerge spon... more Self-organization describes the evolution process of complex structures where systems emerge spontaneously, driven internally by variations of the system itself. Self-organization to the critical state is manifested by scale-free behavior across many orders of magnitude (Bak et al. 1987, Bak 1996, Solé et al. 1999). Spatial scale-free behavior implies fractal properties and is quantified by the fractal dimension. Temporal scale-free behavior is evident in power spectra of fluctuations that obey power laws. Self-organized criticality is a universal phenomenon that likely produces some of the fractals and power laws observed in nature. We investigated the historical landscape of southern Wisconsin (USA) (60,000 km 2) for self-organization and complexity. The landscape is patterned into prairies, savannas, and open and closed forests, using data from the United States General Land Office Surveys that were conducted during the 19th century, at a time prior to Euro-American settlement. We applied a two-dimensional cellular automaton model with one adjustable parameter. Model evolution replaces a cell that dies at random times by a cell chosen randomly from within a circular radius r, where r typically takes values between 1 (local) and 10 units (regional). Cluster probability is used to measure the degree of organization. The model landscape self-organizes to a realistic critical state if neighborhoods of intermediate size (r= 3) are chosen, indicating that (a) no particular time or space scale for the clusters is singled out, i.e. the spatial dependence is fractal, and temporal fluctuations in the cluster probability exhibit power laws; (b) a simple model suffices to replicate the landscape pattern resulting from complex spatial and temporal interactions. Measures of comparison between the observed and the simulated landscape show good agreement: fractal dimensions for simulated (1.6) and observed landscapes (1.64), cluster probabilities for simulated (32.3%) and observed (32.6%) landscapes, and algorithmic complexity for simulated (6792 bytes) and observed (6205 bytes) landscapes. The results are robust towards variation of initial and boundary conditions as well as perturbations.

Landscape Ecology, 2007
Our research illustrates how a landscape mosaic changes in association with a mixed naturalanthro... more Our research illustrates how a landscape mosaic changes in association with a mixed naturalanthropogenic disturbance history. Our study area is the Northwest Wisconsin (USA) Sand Plain (NWSP), a region with a rich disturbance history including fire, insects and clearcut forestry. We integrated historic airphotos from 1938, 1960, 1980 and 1998 within a GIS to describe change among four landcover classes describing a canopy-closure gradient: closed forests, woodlands, savannas and ''open barrens''. Our work addresses two literature needs: empirical studies of mixed-disturbance landscapes, and nonforest habitats within a forest matrix. Our analysis shows that: the area of open barrens fluctuated, woodlands and savannas declined severely and closed forests increased through time. Falling median patch sizes and other landscape metrics suggest that the woodlands are becoming more fragmented. The landcover transitions driving this change vary according to time and place. The dominant transitions are toward closed forests from all classes, and transitions toward open barrens are also consistently important. The woodlands, savannas and open barrens habitats are mostly comprised of transient patches, persisting for less than 20 years. This contrasts with closed forests that often persist for 40 plus years. These changes are consistent with the disturbance regime that is shifting from fire-to forestry-dominance. Our results show a trend towards landscape simplification, manifest as losses of intermediate-density habitats (woodland and savanna) and shrinking patch sizes. The transient nature of the nonforest habitats shows that disturbance resulting in total or partial canopy removal will be vital for their conservation at a landscape scale.

Landscape Ecology, 2004
We used the LANDIS disturbance and succession model to study the effects of six alternative veget... more We used the LANDIS disturbance and succession model to study the effects of six alternative vegetation management scenarios on forest succession and the subsequent risk of canopy fire on a 2791 km 2 landscape in northern Wisconsin, USA. The study area is a mix of fire-prone and fire-resistant land types. The alternatives vary the spatial distribution of vegetation management activities to meet objectives primarily related to forest composition and recreation. The model simulates the spatial dynamics of differential reproduction, dispersal, and succession patterns using the vital attributes of species as they are influenced by the abiotic environment and disturbance. We simulated 50 replicates of each management alternative and recorded the presence of species age cohorts capable of sustaining canopy fire and the occurrence of fire over 250 years. We combined these maps of fuel and fire to map the probability of canopy fires across replicates for each alternative. Canopy fire probability varied considerably by land type. There was also a subtle, but significant effect of management alternative, and there was a significant interaction between land type and management alternative. The species associated with high-risk fuels ͑conifers͒ tend to be favored by management alternatives with more disturbances, whereas low disturbance levels favor low-risk northern hardwood systems dominated by sugar maple. The effect of management alternative on fire risk to individual human communities was not consistent across the landscape. Our results highlight the value of the LANDIS model for identifying specific locations where interacting factors of land type and management strategy increase fire risk.

Journal of Biogeography, 2005
Aim Predictions of ecosystem responses to climate warming are often made using gap models, which... more Aim Predictions of ecosystem responses to climate warming are often made using gap models, which are among the most effective tools for assessing the effects of climate change on forest composition and structure. Gap models do not generally account for broad‐scale effects such as the spatial configuration of the simulated forest ecosystems, disturbance, and seed dispersal, which extend beyond the simulation plots and are important under changing climates. In this study we incorporate the broad‐scale spatial effects (spatial configurations of the simulated forest ecosystems, seed dispersal and fire disturbance) in simulating forest responses to climate warming. We chose the Changbai Natural Reserve in China as our study area. Our aim is to reveal the spatial effects in simulating forest responses to climate warming and make new predictions by incorporating these effects in the Changbai Natural Reserve.Location Changbai Natural Reserve, north‐eastern China.Method We used a coupled ...

Global Change Biology, 2005
In the coming century, forecast climate changes caused by increasing greenhouse gases may produce... more In the coming century, forecast climate changes caused by increasing greenhouse gases may produce dramatic shifts in tree species distributions and the rates at which individual tree species sequester carbon or release carbon back to the atmosphere. The species composition and carbon storage capacity of northern Wisconsin (USA) forests are expected to change significantly as a result. Projected temperature changes are relatively large (up to a 5.8°C increase in mean annual temperature) and these forests encompass a broad ecotone that may be particularly sensitive to climate change. Our objective was to estimate the combined effects of climate change, common disturbances, and species migrations on regional forests using spatially interactive simulations. Multiple scenarios were simulated for 200 years to estimate aboveground live biomass and tree species composition. We used a spatially interactive forest landscape model (LANDIS‐II) that includes individual tree species, biomass accu...

Forest Ecology and Management, 2007
A growing trend in public forest management is the inclusion of the public in the decision making... more A growing trend in public forest management is the inclusion of the public in the decision making process. Visual representations of the management process can assist in conveying complex management treatments. A second trend has been the promotion of biological diversity as a management objective. Ecosystem managers and restoration ecologists are using pre-settlement landscape patterns and forest conditions as a reference point to encourage the recovery of rare or extirpated species and habitat types. The problem is that information about pre-settlement conditions is limited. Our research goal was to visualize pre-settlement forests in Wisconsin and compare them with current forest conditions. Presettlement vegetation conditions were derived from computerized U.S. Public Land Survey records. Current forest conditions were derived from USDA Forest Service Forest Inventory and Analysis data. We used World Construction Set software (3D Nature, LLC) for the visualizations. Our results focus on ecosystems that are (a) still widespread but altered in structure and species composition (northern hardwoods communities) or (b) greatly diminished in extent (pine barrens communities). We found there are substantial visual differences between current and pre-settlement forests, most notably in species composition, density, and stand structural complexity. Our results highlight the potential of computer visualization as a tool to aid forest managers and restoration ecologists.

Forest Ecology and Management, 2006
Forest management shapes landscape patterns, and these patterns often differ significantly from t... more Forest management shapes landscape patterns, and these patterns often differ significantly from those typical for natural disturbance regimes. This may affect wildlife habitat and other aspects of ecosystem function. Our objective was to examine the effects of different forest management decisions on landscape pattern in a fire adapted ecosystem. We used a factorial design experiment in LANDIS (a forest landscape simulation model) to test the effects of: (a) cut unit size, (b) minimum harvest age and (c) target species for management. Our study area was the Pine Barrens of northwest Wisconsin, an area where fire suppression has caused a lack of large open areas important for wildlife. Our results show that all three management choices under investigation (cut unit size, minimum harvest age and target species for management) have strong effects on forest composition and landscape patterns. Cut unit size is the most important factor influencing landscape pattern, followed by target species for management (either jack pine or red pine) and then minimum harvest age. Open areas are more abundant, and their average size is larger, when cut units are larger, target species is jack pine, and minimum harvest age is lower. Such information can assist forest managers to relate stand level management decision to landscape patterns.

Forest Ecology and Management, 2008
Changes in land use patterns in and around forests, including rural development and road building... more Changes in land use patterns in and around forests, including rural development and road building, have occurred throughout the United States and are accelerating in many areas. As a result, there have been significant departures from 'natural' or pre-settlement disturbance regimes. Altered disturbance regimes can shift composition and dominance in tree species communities, potentially affecting ecosystem functioning. We examined the potential consequences of various forest management practices and forest fragmentation on tree community composition. Both forest management and fragmentation are changing as land use changes within the New Jersey Pine Barrens (NJPB). The NJPB has and is continuing to experience rapid rural development and urbanization that are altering the types, frequency, and intensity of forest management, and are increasing forest fragmentation. In the NJPB, the size and frequency of wildfires have declined and the use of prescribed fires is limited to a small portion of the landscape. In addition, the expansion of roads and decline in total forested area-two common measures of fragmentation-may impede the ability of tree species to colonize available habitat. To assess the consequences of fire management and fragmentation on fire regimes and forest communities, we simulated forest landscape change using LANDIS-II, a stochastic, spatially dynamic forest succession model that simulates the growth of tree species cohorts (defined by species and age), dispersal and colonization, and mortality. Simulated fires are sensitive to fuel loads and fuel load continuity. We constructed scenarios to mimic the pre-colonial contiguous landscape with an estimated pre-colonial fire regime; scenarios of the current day landscape with current and potential fire management; and scenarios designed to highlight the effects of fragmentation. Our simulations indicate that relative to the pre-colonial landscape and fire regime, the landscape is changing from a pine-dominated to an oakdominated state. However, within areas where prescribed burning remains a viable management option, a doubling of the mean annual area that is managed with prescribed burns may substantially push the system back towards pre-colonial conditions, although oaks will continue to retain a higher than pre-colonial dominance. Our results also indicate that aside from a reduction in the potential fire sizes, fragmentation does not appear to substantially alter forest successional dynamics. In summary, our simulations estimate the departure from pre-colonial conditions and indicate the potential for a limited restoration of these conditions.

Forest Ecology and Management, 2002
We examined tree species responses under forest harvesting and an increased fire disturbance scen... more We examined tree species responses under forest harvesting and an increased fire disturbance scenario due to climate warming in northern Wisconsin where northern hardwood and boreal forests are currently predominant. Individual species response at the ecosystem scale was simulated with a gap model, which integrates soil, climate and species data, stratified by ecoregions. Such responses were quantified as species establishment coefficients. These coefficients were used to parameterize a spatially explicit landscape model, LANDIS. Species response to climate warming at the landscape scale was simulated with LANDIS, which integrates ecosystem dynamics with spatial processes including seed dispersal, fire disturbance, and forest harvesting. Under a 5 8C annual temperature increase predicted by global climate models (GCM), our simulation results suggest that significant change in species composition and abundance could occur in the two ecoregions in the study area. In the glacial lake plain (lakeshore) ecoregion under warming conditions, boreal and northern hardwood species such as red oak, sugar maple, white pine, balsam fir, paper birch, yellow birch, and aspen decline gradually during and after climate warming. Southern species such as white ash, hickory, bur oak, black oak, and white oak, which are present in minor amounts before the warming, increase in abundance on the landscape. The transition of the northern hardwood and boreal forest to one dominated by southern species occurs around year 200. In the sand barrens ecoregion under warming conditions, red pine initially benefits from the decline of other northern hardwood species, and its abundance quickly increases. However, red pine and jack pine as well as new southern species are unable to reproduce, and the ecoregion could transform into a region with only grass and shrub species around 250 years under warming climate. Increased fire frequency can accelerate the decline of shadetolerant species such as balsam fir and sugar maple and accelerate the northward migration of southern species. Forest harvesting accelerated the decline of northern hardwood and boreal tree species. This is especially obvious on the barrens ecoregion, where the intensive cutting regime contributed to the decline of red pine and jack pine already under stressed environments. Forest managers may instead consider a conservative cutting plan or protective management scenarios with limited forest harvesting. This could prolong the transformation of the barrens into prairie from one-half to one tree life cycle.

Environmental Modelling & Software, 2002
Forest management issues are increasingly required to be addressed in a spatial context, which ha... more Forest management issues are increasingly required to be addressed in a spatial context, which has led to the development of spatially explicit forest landscape models. The numerous processes, complex spatial interactions, and diverse applications in spatial modeling make the development of forest landscape models difficult for any single research group. New developments in componentbased modeling approaches provide a viable solution. Component-based modeling breaks a monolithic model into small, interchangeable, and binary components. They have these advantages compared to the traditional modeling work: 1) developing a component is a much smaller task than developing a whole model, 2) a component can be developed using most programming languages, since the interface format is binary, and 3) new components can replace the existing ones under the same model framework; this reduces the duplication and allows the modeling community to focus resources on the common products, and to compare results. In this paper, we explore the design of a spatially explicit forest landscape model in a component-based modeling framework, based on our work on object-oriented forest landscape modeling. We examine the representation of the major components and the interactions between them. Our goal is to facilitate the use of the component-based modeling approach at the early stage of spatially explicit landscape modeling.

Environmental Conservation, 2013
SUMMARYSpatial simulation may be used to model the potential effects of current biodiversity appr... more SUMMARYSpatial simulation may be used to model the potential effects of current biodiversity approaches on future habitat modification under differing climate change scenarios. To illustrate the approach, spatial simulation models, including landscape-level forest dynamics, were developed for a semi-natural grassland of conservation concern in a southern Italian protected area, which was exposed to woody vegetation encroachment. A forest landscape dynamics simulator (LANDIS-II) under conditions of climate change, current fire and alternative management regimes was used to develop scenario maps. Landscape pattern metrics provided data on fragmentation and habitat quality degradation, and quantified the spatial spread of different tree species within grassland habitats. The models indicated that approximately one-third of the grassland area would be impacted by loss, fragmentation and degradation in the next 150 years. Differing forest management regimes appear to influence the type o...
Ecosystems, 2012
Your article is protected by copyright and all rights are held exclusively by Springer Science +B... more Your article is protected by copyright and all rights are held exclusively by Springer Science +Business Media New York. This e-offprint is for personal use only and shall not be selfarchived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com".

Ecosystems, 1999
The study of forest landscape change requires an understanding of the complex interactions of bot... more The study of forest landscape change requires an understanding of the complex interactions of both spatial and temporal factors. Traditionally, forest gap models have been used to simulate change on small and independent plots. While gap models are useful in examining forest ecological dynamics across temporal scales, large, spatial processes, such as seed dispersal, cannot be realistically simulated across large landscapes. To simulate seed dispersal, spatially explicit landscape models that track individual species distribution are needed. We used such a model, LANDIS, to illustrate the implications of seed dispersal for simulating forest landscape change. On an artificial open landscape with a uniform environment, circular-shaped tree species establishment patterns resulted from the simulations, with areas near seed sources more densely covered than areas further from seed sources. Because LANDIS simulates at 10-y time steps, this pattern reflects an integration of various possible dispersal shapes and establishment that are caused by the annual variations in climate and other environmental variables. On real landscapes, these patterns driven only by species dispersal radii are obscured by other factors, such as species competition, disturbance, and landscape structure. To further demonstrate the effects of seed dispersal, we chose a fairly disturbed and fragmented forest landscape (approximately 500,000 ha) in northern Wisconsin. We compared the simulation results of a map with tree species (seed source locations) realistically parameterized (the real scenario) against a randomly parameterized species map (the random scenario). Differences in the initial seed source distribution lead to different simulation results of species abundance with species abundance starting at identical levels under the two scenarios. This is particularly true for the first half of the model run (0-250 y). Under the random scenario, infrequently occurring and shade tolerant species tend to be overestimated, while midabundant and midshade tolerant species tend to be underestimated. The over-and underestimation of species abundance diminish when examining longterm (500 y) landscape dynamics, because stochastic factors, such as fire, tend to make the landscapes under both scenarios converge. However, differences in spatial patterns, and especially species age-cohort distributions, can persist under the two scenarios for several hundred years.

Ecology, 2005
Within the northern Great Lakes region, mesoscale (10s to 100s of km2) forest patterning is drive... more Within the northern Great Lakes region, mesoscale (10s to 100s of km2) forest patterning is driven by disturbance dynamics. Using original Public Land Survey (PLS) records in northern Wisconsin, USA, we study spatial patterns of wind and fire disturbances during the pre-Euroamerican settlement period (ca. 1850). Our goals were: (1) to determine how effectively wind and fire disturbance can be reconstructed from the PLS, (2) to assess the roles of wind and fire in shaping vege~ation patterns, (3) to evaluate landscape to regional controls of wind and fire regimes, and (4) to assess the potential for interactions between these disturbances. Our analyses indicate that only relatively intense fire and wind disturbance can be reliably detected from the PLS (62-68% canopy removal). Heavy windthrow was more prevalent than fire disturbance in presettlement forests, and wind-disturbed patches were comparatively smaller and tnore complex in shape. Disturbance rotation periods ranged between 450 and 10 500 years for heavy windthrow and between 700 and 93 000 years for stand-replacing fire. Occurrences of wind and fire disturbance were related to geographic province and to regional soil patterns: analysis further suggests a negative interaction between the two disturbance types. Given that severe wind disturbance was infrequent, mature to old forests of late-successional species dominated much of pre-Euroamerican northern Wisconsin, but wind disturbances may have allowed regional persistence of less shade-tolerant species, such as Betula alleghaniensis. Pine-dominated vegetation was limited to regions with more frequent fire, but frequencies of stand-replacing fire derived from survey records were insufficient to maintain these successional vegetation types; we suggest that frequent surface fires, not recorded in the PLS, along with infrequent stand-replacing fire, maintained these vegetation types. The extensive nature of the PLS provides a powerful baseline for addressing changes in forest conditions and disturbance regimes associated with climate and land use for both the present and more distant past. Such baselines are informative in discussions of historical variability and restoration silviculture.

Ecology, 1999
Understanding disturbance and recovery of forest landscapes is a challenge because of complex int... more Understanding disturbance and recovery of forest landscapes is a challenge because of complex interactions over a range of temporal and spatial scales. Landscape _ O _, simulation models offer an approach to studying such systems at broad scales. Fire can be _ _ _ I:_ simulated spatially using mechanistic or stochastic approaches. We describe the fire module f0 to ;-,in a spatially explicit, stochastic model of forest hmdscape dynamics (LANDIS) that in-0_ el" O 6'1 l'a-_ corporates fire, windthrow, and harvest disturbance with species-level succession. A sto-_. _ clchastlc approach is suited to forest landscape models that are designed to simulate patterns _ over large spatial and time domains and are not used deterministically to predict individual !:_m _ events, _ _ We used the model to examine how disturbance regimes and species dynamics interact _.ft_ _ _ across a large (500000 ha), heterogeneous landscape in northern Wisconsin, USA, with six land types having different species environments, and fire disturbance return intervals _ _. _" varying from 200 to 1000 yr. The model shows that there are feedbacks over time between species, disturbance, and environment, resulting in the re-emergence of patterns that char-' _ _" acterized the landscape before extensive alteration. Landscape equilibrium of species corn-_ _, _"°p osition and age-class structure develops at three scales from the initial, disturbed landscape. ._ t.o resilience; succession; verification and calibration.
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Papers by David Mladenoff