Modeling the effects of forest harvesting on landscape structure and the spatial distribution of cowbird brood parasitism

Timber harvesting affects both composition and structure of the landscape and has important consequences for organisms using forest habitats. A timber harvest allocation model was constructed that allows the input of specific rules to allocate forest stands for clearcutting to generate landscape patterns reflecting the look and feel of managed landscapes. Various harvest strategies were simulated on four 237 km² study areas in Indiana, USA. For each study area, the model was applied to simulate 80 years of management activity. The resulting landscape spatial patterns were quantified using a suite of landscape pattern metrics and plotted as a function of mean harvest size and total area of forest harvested per decade to produce response surfaces. When the mean clearcut size was 1 ha, the area of forest interior remaining on the landscape was dramatically reduced and the amount of forest edge on the landscape increased dramatically. The potential consequences of the patterns produced by the model were assessed for a generalized neotropical migrant forest bird using a GIS model that generates maps showing the spatial distribution of the relative vulnerability of forest birds to brood parasitism by brown-headed cowbirds. The model incorporates the location and relative quality of cowbird feeding sites, and the relation between parasitism rates and distance of forest from edge. The response surface relating mean harvest size and total area harvested to the mean value of vulnerability to cowbird brood parasitism had a shape similar to the response surfaces showing forest edge. The results of our study suggest that it is more difficult to maintain large contiguous blocks of undisturbed forest interior when harvests are small and dispersed, especially when producing high timber volumes is a management goal. The application of the cowbird model to landscapes managed under different strategies could help managers in deciding where harvest activity will produce the least negative impact on breeding forest birds.     

Human influence on the abundance and connectivity of high-risk fuels in mixed forests of northern Wisconsin,USA

Though fire is considered a natural disturbance, humans heavily influence modern wildfire regimes. Humans influence fires both directly, by igniting and suppressing fires, and indirectly, by either altering vegetation, climate, or both. We used the LANDIS disturbance and succession model to compare the relative importance of a direct human influence (suppression of low intensity surface fires) with an indirect human influence (timber harvest) on the long-term abundance and connectivity of high-risk fuel in a 2791 km² landscape characterized by a mixture of northern hardwood and boreal tree species in northern Wisconsin. High risk fuels were defined as a combination of sites recently disturbed by wind and sites containing conifer species/cohorts that might serve as ladder fuel to carry a surface fire into the canopy. Two levels of surface fire suppression (high/current and low) and three harvest alternatives (no harvest, hardwood emphasis, and pine emphasis) were compared in a 2×3 factorial design using 5 replicated simulations per treatment combination over a 250-year period. Multivariate analysis of variance indicated that the landscape pattern of high-risk fuel (proportion of landscape, mean patch size, nearest neighbor distance, and juxtaposition with non fuel sites) was significantly influenced by both surface fire suppression and by forest harvest (p > 0.0001). However, the two human influences also interacted with each other (p < 0.001), because fire suppression was less likely to influence fuel connectivity when harvest disturbance was simultaneously applied. Temporal patterns observed for each of seven conifer species indicated that disturbances by either fire or harvest encouraged the establishment of moderately shade-tolerant conifer species by disturbing the dominant shade tolerant competitor, sugar maple. Our results conflict with commonly reported relationships between fire suppression and fire risk observed within the interior west of the United States, and illustrate the importance of understanding key interactions between natural disturbance, human disturbance, and successional responses to these disturbance types that will eventually dictate future fire risk.This revised version was published online in May 2005 with corrections to the Cover Date.     

Creating landscape patterns by forest cutting: Ecological consequences and principles

Landscape structural characteristics, such as patch size, edge length, and configuration, are altered markedly when management regimes are imposed on primeval landscapes. The ecological consequences of clearcutting patterns were explored by using a model of the dispersed patch or checkerboard system currently practiced on federal forest lands in the western United States. Thresholds in landscape structure were observed on a gradient of percentages of landscape cutover. Probability of disturbance,e.g., wildfire and windthrow, and biotic components,e.g., species diversity and game populations, are highly sensitive to these structural changes. Altering the spatial configuration and size of clearcuts provides an opportunity to create alternative landscapes that differ significantly in their ecological characteristics. Both ecosystem and heterogeneous landscape perspectives are critical in resource management.     

Disturbance propagation by bark beetles as an episodic landscape phenomenon

Landscapes are the resultant of ecological processes and events operating on many different space-time scales. Large scale disturbance is recognized as a major influence on landscape patterns, but the impact of small scale events is often overlooked. We develop an hierarchical framework to relate lightning and bark beetle population dynamics to the southern pine forest landscape using the concepts of disturbance propagation and amplification. The low level lightning disturbance can be propagated to the landscape level when weather and forest stand structure facilitate bark beetle epidemics. We identify epidemics as biotically-driven episodes that alter landscape structure. The concept of the landscape as the spatial dimension of these episodes is represented in a conceptual model linking insect-host and landscape mosaic interactions.     

Three objectives of historical ecology: the case of litter collecting in Central European forests

Short- and long-term patterns of net ecosystem carbon balance (NECB) for small, relatively uniform forest stands have been examined in detail, but the same is not true for landscapes, especially those with heterogeneous disturbance histories. In this paper, we explore the effect of two contrasting types of disturbances (i.e., fire and tree harvest) on landscape level NECB by using an ecosystem process model that explicitly accounts for changes in carbon (C) stores as a function of disturbance regimes. The latter were defined by the average disturbance interval, the regularity of the disturbance interval (i.e., random, based on a Poisson frequency distribution, or regular), the amount of C removed by the disturbance (i.e., severity), and the relative abundance of stands in the landscape with unique disturbance histories. We used the model to create over 300 hypothetical landscapes, each with a different disturbance regime, by simulating up to 200 unique stand histories and averaging their total C stores. Mean NECB and its year-to-year variability was computed by calculating the difference in mean total C stores from one year to the next. Results indicated that landscape C stores were higher for random than for regular disturbance intervals, and increased as the mean disturbance interval increased and as the disturbance severity decreased. For example, C storage was reduced by 58% when the fire interval was shortened from 250 years to 100 years. Average landscape NECB was not significantly different than zero for any of the simulated landscapes. Year-to-year variability in landscape NECB, however, was related to the landscape disturbance regime; increasing with disturbance severity and frequency, and higher for random versus regular disturbance intervals. We conclude that landscape C stores of forest systems can be predicted using the concept of disturbance regimes, a result that may be a useful for adjusting estimates of C storage to broad scales that are solely based on physiological processes.     

The influence of forest harvesting on landscape spatial patterns and old-growth-forest fragmentation in southeast British Columbia

Habitat fragmentation is considered one of the major conservation issues of recent decades. We tested predictions of landscape patterns in a 352,253-ha managed forest area in southeast British Columbia. We did this by focussing on forest fragmentation concerns among old-growth, harvest, and wildfire patches in 44 delineated landscapes using patch indices as measures of landscape pattern. We found no significant association between amount of harvesting and 15 old-growth patch indices. Comparisons among patch types revealed that amounts and spatial patterns of harvest patches differed little from amounts and spatial patterns of old-growth patches in control landscapes. Variability indices revealed similar variability between harvest patches and old-growth patches, and more variability between harvest patches and wildfire patches. Little of the evidence gathered in this study supported predictions of fragmentation of old-growth spatial patterns, or predicted differences between harvest spatial patterns and more naturally occurring spatial patterns. We suggest these results could be due to the relatively small amounts of harvesting and old-growth forest in these landscapes, and therefore habitat amount may be a more important factor than spatial configuration of patches in these landscapes.     

Watershed analysis of forest fragmentation by clearcuts and roads in a Wyoming forest

Remotely sensed data and a Geographic Information System were used to compare the effects of clearcutting and road-building on the landscape pattern of the Bighorn National Forest, in north-central Wyoming. Landscape patterns were quantified for each of 12 watersheds on a series of four maps that differed only in the degree of clearcutting and road density. We analyzed several landscape pattern metrics for the landscape as a whole and for the lodgepole pine and spruce/fir cover classes across these maps, and determined the relative effects of clearcutting and road building on the pattern of each watershed. At both the landscape- and cover class-scales, clearcutting and road building resulted in increased fragmentation as represented by a distinct suite of landscape structural changes. Patch core area and mean patch size decreased, and edge density and patch density increased as a result of clearcuts and roads. Clearcuts and roads simplified patch shapes at the landscape scale, but increased the complexity of lodgepole pine patches. Roads appeared to be a more significant agent of change than clearcuts, and roads which were more evenly distributed across a watershed had a greater effect on landscape pattern than did those which were densely clustered. Examining individual watersheds allows for the comparison of fragmentation among watersheds, as well as across the landscape as a whole. Similar studies of landscape structure in other National Forests and on other public lands may help to identify and prevent further fragmentation of these areas.     

Cumulative effects of roads and logging on landscape structure in the San Juan Mountains,Colorado (USA)

In the southern Rocky Mountains of temperate North America, the effects of Euro-American activities on disturbance regimes and landscape patterns have been less ubiquitous and less straightforward in high-elevation landscapes than in low-elevation landscapes. Despite apparently little change in the natural disturbance regime, there is increasing concern that forest management activities related mainly to timber harvest and to the extensive network of roads constructed to support timber harvest, fire control, and recreation since the late 1800s have altered disturbance regimes and landscape structure. We investigated the magnitude of change in landscape structure resulting from roads and logging since the onset of timber harvest activities in 1950. We found limited evidence for significant impacts in our study area when all lands within the landscape were considered. The relatively minor changes we observed reflected the vast buffering capacity of the large proportion of lands managed for purposes other than timber (e.g., wilderness). Significant changes in landscape structure and fragmentation of mature forest were, however, evident on lands designated as suitable timberlands. Roughly half of the mature coniferous forest was converted to young stands; mean patch size and core area declined by 40% and 25%, respectively, and contrast-weighted edge density increased 2- to 3-fold. Overall, roads had a greater impact on landscape structure than logging in our study area. Indeed, the 3-fold increase in road density between 1950–1993 accounted for most of the changes in landscape configuration associated with mean patch size, edge density, and core area. The extent of area evaluated and the period over which change was evaluated had a large impact on the magnitude of change detected and our conclusions regarding the ecological significance of those changes. Specifically, the cumulative impact on landscape structure was negligible over a 10-year period, but was notable over a 40-year period. In addition, the magnitude of change in landscape structure between 1950–1993 varied as a function of landscape extent. At the scale of the 228000 ha landscape, change in landscape structure was trivial, suggesting that the landscape was capable of fully incorporating the disturbances with minimal impact. However, at intermediate scales of 1000–10000 ha landscapes, change in landscape structure was quite evident, suggesting that there may be an optimal range of scales for detecting changes in landscape structure within the study area.     

Landscape-scale characteristics of forest tornado damage in mountainous terrain

Context

Landscape patterns created by natural disturbance such as windstorms can affect forest regeneration, carbon cycling, and other ecological processes.

Objectives

We develop a method for remotely measuring tornado damage severity and describe landscape-scale patterns of tornado damage. We examine the extent and distribution of damage severity and gaps created by tornadoes, and examine how topographic variation can influence tornado damage severity.

Methods

Focusing on two April 2011 tornadoes that struck the Chattahoochee National Forest (CNF) in Georgia and the Great Smoky Mountains (GSM) in Tennessee, we used supervised classification of aerial photographs to map damage severity. We report the extent and distribution of damage severity from each track and characterize patterns of damage using FragStats. Using topographic overlays, we test hypotheses regarding how physiographic features such as valleys and ridges affect tornado damage severity.

Results

Tornado damage severity estimates were significantly correlated with ground-truth measurements. The 64-km CNF track damaged 1712 ha (>25 % severity), while the 26-km GSM track damaged 1407 ha. Tornado damage severity was extremely variable and frequency of gap sizes drastically decreased with size, with many small gaps and few very large gaps, consistent with other types of wind damage. Damage severity declined as tornadoes ascended ridges and increased as they descended ridges. This effect was more consistent on shallow slopes relative to steeper slopes.

Conclusions

This study outlines an objective methodology for remotely characterizing tornado damage severity. The results from this study fill an important gap in ecological understanding of the spatial components of the forest tornado disturbance regime.

     

Landscape Fragmentation and Ice Storm Damage in Eastern Ontario Forests

With return times between 20 and 100 years, ice storms are a primary disturbance type for temperate forests of eastern North America. Many studies have been conducted at the forest patch and plot scales to examine relations between damage and variables describing site, composition and structure. This paper presents results from a landscape scale study of fragmentation relations with damage in eastern Ontario forests. Data previously collected for two independent and spatially non-overlapping patch level damage studies were used. A Generalized Linear Model (GLM) was used to analyse relations between damage and fragmentation metrics representing patch isolation, edge density, and the relative size and distribution of patches in the landscape. The metrics were applied using spatial extents of 1 × 1 km and 4 × 4 km, following analyses of the variability of numbers of patches and of the lacunarity of forest patterns over a range of extents. The results showed that patch isolation, as measured by the mean Euclidean distance between patches (ENN) was significantly related to damage.     

Cumulative patterns of logging and fire (1940–2009): consequences on the structure of the eastern Canadian boreal forest

Context

Although logging has affected circumboreal forest dynamics for nearly a century, very few studies have reconstructed its influence on landscape structure at the subcontinental scale.

Objectives

This study aims to document spatiotemporal patterns of logging and fire since the introduction of logging in the early twentieth-century, and to evaluate the effects of these disturbances on landscape structure.

Methods

We used historical (1940–2009) logging and fire maps to document disturbance patterns across a 195,000-km² boreal forest landscape of eastern Canada. We produced multitemporal (1970s–2010s) mosaics providing land cover status using Landsat imagery.

Results

Logging significantly increased the rate of disturbance (+74 %) in the study area. The area affected by logging increased linearly with time resulting in a significant rejuvenation of the landscape along the harvesting pattern (south–north progression). From 1940 to 2009, fire was the dominant disturbance and showed a more random spatial distribution than logging. The recent increase of fire influence and the expansion of the proportion of area classified as unproductive terrestrial land suggest that regeneration failures occurred.

Conclusions

This study reveals how logging has modified the disturbances dynamics, following the progression of the logging frontier. Future management practices should aim for a dispersed spatial distribution of harvests to generate landscape structures that are closer to natural conditions, in line with ecosystem-based management. The challenges of defining sustainable practices will remain complex with the predicted increase in fire frequency, since this factor, in combination with logging, can alter both the structure and potentially the resilience of boreal forest.

     

Spatial pattern analysis of pre- and post-hurricane forest canopy structure in North Carolina,USA

Existing spatial patterns of a forest are in part a product of its disturbance history. Using laser altimetry and field measures of canopy top height to represent pre- and post-hurricane canopy topography, respectively, we measured changes in spatial patterns of stand structure of a United States southern mixed coniferous-deciduous for est. Autocorrelative and fractal properties were measured in this opportunistic study to quantify changes in canopy architecture along twelve, 190-250 m transects that were subjected to moderate to high levels of wind disturbance. Prior to the hurricane, canopy heights were autocorrelated at scales <40 m with an average fractal dimension of 1.71. After the disturbance, autocorrelation disappeared; the average fractal dimension rose to 1.94. This shift towards spatial randomness illustrates part of the cyclical nature of ecosystem development. It shows how a catastrophic collapse of biomass accumulation corresponds to a decrease in ecosystem organization across a landscape. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spatial patterns of bird community similarity: bird responses to landscape composition and configuration in the Atlantic forest

Studies dealing with community similarity are necessary to understand large scale ecological processes causing biodiversity loss and to improve landscape and regional planning. Here, we study landscape variables influencing patterns of community similarity in fragmented and continuous forest landscapes in the Atlantic forest of South America, isolating the effects of forest loss, fragmentation and patterns of land use. Using a grid design, we surveyed birds in 41 square cells of 100 km² using the point count method. We used multivariate, regression analyses and lagged predictor autoregressive models to examine the relative influence of landscape variables on community similarity. Forest cover was the primary variable explaining patterns of bird community similarity. Similarity showed a sudden decline between 20 and 40% of forest cover. Patterns of land use had a second order effect; native bird communities were less affected by forest loss in landscapes dominated by tree plantations (the most suitable habitat for native species) than in landscapes dominated by annual crops or cattle pastures. The effects of fragmentation were inconclusive. The trade-off between local extinctions and the invasion of extra-regional species using recently created habitats is probably the mechanism generating the observed patterns of community similarity. Limiting forest loss to 30–40% of the landscape cover and improving the suitability of human-modified habitats will contribute to maintain the structure and composition of the native forest bird community in the Atlantic forest.     

Developing alternative forest cutting patterns: A simulation approach

This study examines effects of different forest cutting patterns on habitat fragmentation in managed forest landscapes. We use computer simulation to conduct experiments in which we examine effects of different cutting patterns, cutting-unit size, and special constraints (e.g., a forest reserve, a stream system, or a road system) on landscape patterns. Fragmentation indices are used to quantify structural changes over the cutting cycle and among different treatments of the experiments. Degree of fragmentation varies greatly among the five cutting patterns used; aggregation of cutting units results in low degree and gradual change of fragmentation. Cutting patterns with larger cutting units and additional landscape constraints also lead to lower degree of fragmentation. Moreover, differences in fragmentation among the treatments are not observed until 30% or 50% of the landscape is cut.     

The protean relationship between boreal forest landscape structure and red squirrel distribution at multiple spatial scales

This paper investigates two fundamental questions in landscape ecology: what influence does landscape context, or the composition of the matrix, have on an animals’ response to landscape structure, and how does this relationship extrapolate between landscapes? We investigate how the distribution of North American red squirrels (Tamiasciurus hudsonicus) in the boreal mixedwood forest is influenced by anthropogenically (forest harvest) and naturally (forest fire) derived landscape structure. We studied the presence and absence of red squirrels over two years in three landscape types: one managed for timber harvest, one recently burned by wildfire, and a third unburned unmanaged landscape. Landscape composition and configuration, measured at several spatial scales, predicted red squirrel’s distribution in all three landscapes, but the significant landscape variables changed across spatial scales, across time, and across landscapes. These findings emphasize the variability in landscape structure/animal distribution relationships, and enforce the need to link pattern-finding studies, such as this one, with searches for the mechanisms behind the observed pattern.     

Spatial variability in soil nitrogen dynamics after prescribed burning in Ohio mixed-oak forests

This study describes the results of the application of a single dormant season prescribed fire to two southern Ohio forest sites for the purposes of restoring the ecosystem functional properties that existed in these sites prior to major human intervention (clearcutting, fire suppression, and atmospheric deposition). Each forest site was composed of three contiguous watershed units, two of which were burned in April of 1996. The forest sites differed in soil pH and available litter mass prior to the fires, and in both sites pH and available inorganic N varied among landscape positions such that inorganic C increased with increasing longterm soil moisture potential (measured as the GIS-derived Integrated Moisture Index [IMI] developed for this region). The fire temperatures at 10 cm above the litter surface were generally 150–300°C, and 29–80% of the litter was consumed, depending on site and landscape position. Soil solution total inorganic N (TIN) present one month after the fires did not differ significantly from that present prior to the fires in either burned or unburned watersheds, but was consistently greater in mesic landscape positions than in more xeric ones. N mineralization potential and organic C content varied both among fires and landscape positions. At the site which burned at higher intensity, soil N mineralization and TIN were both decreased by fire. At the less intensely burned site, fire resulted in increased TIN in the soils from the more xeric landscape position, and greater soil organic C in soils from the intermediate soil moisture areas. Path analysis produced models for fire-induced changes in C and N dynamics capable of explaining 26–69% of the observed variation using combinations of landscape and fire behavior. Losses of N to volatilization from these single fires were generally < 1 kg N/ha, and thus could not be expected to ameliorate the effects of atmospheric N deposition in these sites.     

Physical influences of landscape on a large-extent ecological disturbance: the northeastern North American ice storm of 1998

The 1998 ice storm was a large-extent ecological disturbance that severely affected the eastern Adirondack forests of northern New York. Ice damage produced widespread breakage of limbs and trunks in susceptible trees. Although ice storms are common within northeastern North American forests, the magnitude and extent of the 1998 storm far exceeded damage caused by typical ice storms in the recent past. While plot and stand-scale ecological impacts of ice storms have received attention insofar as tree species vulnerability, stand age susceptibility, and microhabitat alterations, larger-extent damage patterns have not been previously evaluated. The normalized difference vegetation index (NDVI) was employed to assess forest vigor and canopy density in atmospherically corrected Landsat Thematic Mapper (TM) satellite imagery of the Adirondacks. Digital change analysis of the baseline forest condition (1990 NDVI data), and the condition encountered in a post-storm image (1998 NDVI data) was conducted. Forest damage was separated from natural variations in canopy reflectance by employing a generalized linear model that incorporated in situ measurements. A robust empirical variogram analysis revealed that locations of tree damage were significantly correlated for distances up to 300 meters. Intensely-damaged forest exhibited greater spatial dependence, but over a smaller distance. Canopy damage was not greater proximate to stream and forest boundaries, and did not follow our hypothesis of decreasing damage with distance from the boundary. Overall, we show that local topography (elevation and aspect), forest composition (deciduous or coniferous), and the meteorological characteristics of the disturbance event acted together to determine the spatial extent of ice storm damage.This revised version was published online in May 2005 with corrections to the Cover Date.     

Using landscape metrics to model source habitat for Neotropical migrants in the midwestern U.S.

Size of a forest patch is a useful predictor of density and reproductive success of Neotropical migratory birds in much of eastern North America. Within these forested landscapes, large forest tracts appear to be sources – fragments in which surpluses of offspring are produced and can potentially colonize new fragments including woodlot sinks where reproduction fails to balance adult mortality. Within agricultural landscapes of the midwestern U.S., where forests are severely fragmented, high levels of brood parasitism by brown-headed cowbirds (Molothrus ater) and intense predation on nests generally result in low reproductive success for Neotropical migrants regardless of forest size. In some midwestern U.S. landscapes, however, the variation in reproductive success among forest fragments suggests that `source' habitat could still exist for Neotropical migrants. We used vegetation, fragment and landscape metrics to develop multivariate models that attempt to explain the variation in abundance and reproductive success of Neotropical migrants nesting in an agricultural landscape in northern Indiana, USA. We produced models that reasonably described the pattern of species richness of Neotropical migrants and the abundance of wood thrushes (Hylocichla mustelina) and several other Neotropical migrant species within 14 forest fragments. In contrast, we were unable to produce useful models of the reproductive success of wood thrushes breeding in the same forest fragments. Our results suggest that (1) abundance patterns of Neotropical migrants are probably influenced by both landscape- and fragment-scale factors; (2) multivariate analyses of Neotropical migrant abundance are not useful in modeling the corresponding patterns of reproductive success; and (3) the location of any remaining `source' habitat for Neotropical migrants breeding within agricultural landscapes in North America will be difficult to predict with indirect measures such as vegetation composition or landscape context. As a result, the potential for developing conservation strategies for Neotropical migrants will be limited without labor-intensive, direct measurements of demographic parameters.     

Historic range of variability in landscape structure in subalpine forests of the Greater Yellowstone Area,USA

A measure of the historic range of variability (HRV) in landscape structure is essential for evaluating current landscape patterns of Rocky Mountain coniferous forests that have been subjected to intensive timber harvest. We used a geographic information system (GIS) and FRAGSTATS to calculate key landscape metrics on two ∼130,000-ha landscapes in the Greater Yellowstone Area, USA: one in Yellowstone National Park (YNP), which has been primarily shaped by natural fires, and a second in the adjacent Targhee National Forest (TNF), which has undergone intensive clearcutting for nearly 30 years. Digital maps of the current and historical landscape in YNP were developed from earlier stand age maps developed by Romme and Despain. Maps of the TNF landscape were adapted from United States Forest Service Resource Information System (RIS) data. Key landscape metrics were calculated at 20-yr intervals for YNP for the period from 1705-1995. These metrics were used to first evaluate the relative effects of small vs. large fire events on landscape structure and were then compared to similar metrics calculated for both pre- and post-harvest landscapes of the TNF. Large fires, such as those that burned in 1988, produced a structurally different landscape than did previous, smaller fires (1705-1985). The total number of patches of all types was higher after 1988 (694 vs. 340-404 before 1988), and mean patch size was reduced by almost half (186 ha vs. 319-379 ha). The amount of unburned forest was less following the 1988 fires (63% vs. 72-90% prior to 1988), yet the number of unburned patches increased by nearly an order of magnitude (230 vs. a maximum of 41 prior to 1988). Total core area and mean core area per patch decreased after 1988 relative to smaller fires (∼73,700 ha vs. 87,000-110,000 ha, and 320 ha vs. 2,123 ha, respectively). Notably, only edge density was similar (17 m ha⁻¹ after 1988) to earlier landscapes (9.8-14.2 m ha⁻¹).Three decades of timber harvesting dramatically altered landscape structure in the TNF. Total number of patches increased threefold (1,481 after harvest vs. 437 before harvest), and mean patch size decreased by ∼70% (91.3 ha vs. 309 ha). None of the post-harvest landscape metrics calculated for the TNF fell within the HRV as defined in YNP, even when the post-1988 landscape was considered. In contrast, pre-harvest TNF landscape metrics were all within, or very nearly within, the HRV for YNP. While reference conditions such as those identified by this study are useful for local and regional landscape evaluation and planning, additional research is necessary to understand the consequences of changes in landscape structure for population, community, ecosystem, and landscape function. This revised version was published online in July 2006 with corrections to the Cover Date.